Zootaxa 3061: 53–66 (2011) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2011 · Magnolia Press ISSN1175-5334(online edition) Description and biological observations on a new species of deepwater symphurine tonguefish (Pleuronectiformes: Cynoglossidae: Symphurus) collected at Volcano–19, Tonga Arc, West Pacific Ocean THOMAS A. MUNROE1,3, JENNIFER TYLER2 & VERENA TUNNICLIFFE2 1National Systematics Laboratory, National Marine Fisheries Service, NOAA, Smithsonian Institution, Post Office Box 37012, National Museum of Natural History, WC–57, MRC–153, Washington, DC 20013–7012, USA. E-mail: [email protected] 2Department of Biology, University of Victoria, PO Box 3080, Victoria, British Columbia, Canada V8W 3N5. E-mail: [email protected]; [email protected] 3Corresponding author. E-mail: [email protected] Abstract Symphurus maculopinnisn. sp., described on a single specimen (USNM 398820; 84.4 mm SL), was collected by a re- motely operated vehicle (ROV) exploring a hydrothermal vent area located at 561 m on Volcano–19, Tonga Arc, West Pacific (24°48.439' S, 177°0.009' W). This new species is distinctive and readily diagnosed from congeners by the follow- ing combination of characters: 1–2–2–2–1 pattern of interdigitation of dorsal proximal pterygiophores and neural spines (ID pattern), 14 caudal-fin rays, 3+6 abdominal vertebrae, 49 total vertebrae, 89 scales in a longitudinal row, 92 dorsal- fin rays, 77 anal-fin rays, blunt squarish snout, thick blind-side lips with conspicuous plicae, and conspicuous ocellated (sometimes partially) spots on posterior dorsal and anal fins. Among Symphurus, only S. ocellatus von Bonde, collected at deepwater locations off East Africa, features a similar ID pattern, 14 caudal-fin rays and spots on the posterior dorsal and anal fins. Symphurus maculopinnis differs distinctly from S. ocellatus in its lower and non-overlapping meristic fea- tures (49 vs. 54–56 total vertebrae; 92 vs. 97–103 dorsal-fin rays; and 77 vs. 85–89 anal-fin rays), its squarish (vs. pointed) snout, and thick, plicated blind-side lower lip (vs. thin, non-plicated blind-side lower lip). Additional specimens (N= 56) ofS. maculopinnis observed and filmed in situ near active venting sites located between ca. 433-561 m on Volcano–19 provide the basis for behavioral and ecological information recorded for the species. Videotapes reveal one individual of S. maculopinnis featuring reversed (dextral) asymmetry from that typical (sinistral) for members of the Cynoglossidae. Specimens with reversed asymmetry are relatively rare in this family and this S. maculopinnis represents only the second known reversed individual among the approximately 42 species of deep-sea (>200 m) Symphurus. Key words: taxonomy, deep-sea flatfish, tongue sole, hydrothermal vents, volcanic arcs, seamounts, reversed asymmetry, species description, ocelli Introduction Exploratory expeditions along the Tonga and Kermadec Volcanic Arcs investigated various features of submarine volcanoes, hydrothermal vents and their associated faunas (e.g. Stoffers et al. 2006). Of particular interest during these expeditions was the discovery of symphurine tonguefishes associated with deepwater hydrothermal vents and other habitats on these undersea volcanoes (Tunnicliffe et al. 2010). One undescribed tonguefish species, Species B, observed on Volcano–19 in the Tonga Arc, was conspicuous in the possession of a single pigmented spot on both the posterior dorsal and anal fins. This pigmentation feature is rare among species of Indo–West Pacific Sym- phurus (Munroe unpubl. data), with only S. ocellatus von Bonde collected from deep waters in the western Indian Ocean having such spots on its dorsal and anal fins. A single specimen (now the holotype) of this undescribed species, collected at 561 m on Volcano–19 during a dive conducted by the ROV ROPOS, provides the basis for formal description of this species. Details of the expe- ditions including physico-chemical data of habitats where this specimen was collected and where other specimens of this species were observed, as well as ecological and behavioral observations for the new species, were provided in Tunnicliffe et al. (2010). Accepted by M.T. Craig: 8 Aug. 2011; published: 18 Oct. 2011 53 Material and methods The holotype (USNM 398820) is deposited in the fish collection of the United States National Museum. Descrip- tion of this species is based on counts and measurements from the holotype, with features of live pigmentation pro- vided by a photograph of the holotype prior to collection (Fig. 1) and photographs of additional specimens filmed in situ on Volcano–19 from a remotely operated vehicle (ROV). Comparative materials for other Indo-Pacific spe- cies of Symphurus are detailed in Munroe (1992), Krabbenhoft and Munroe (2003), Munroe (2006), Munroe and Hashimoto (2008), and Lee et al. (2009a). FIGURE 1. Holotype of Symphurus maculopinnis,n. sp. (now USNM 398820), photographed in situ just prior to collection by ROVROPOS at 561 m on Volcano–19, Tonga Arc, West Pacific. Manipulator is holding a scoop net. Methods for counts and measurements and general terminology for systematics of tonguefishes follow those of Munroe (1998). Terminology for interdigitation patterns of proximal dorsal pterygiophores and neural spines (ID pattern) follow that of Munroe (1992). ID patterns, fin ray and vertebral counts were made from radiographs. Standard length (SL) and head length (HL) are used throughout. Measurements were made to the nearest 0.1 mm using either dial calipers or a dissecting stereo microscope fitted with a calibrated ocular micrometer. Morpho- metric features are expressed either as measurements in percentages of standard length, or percentages of head length. Pigmentation of alcohol-preserved fish is based on the holotype, which was originally fixed in 95% ethanol and then transferred and stored in 75% ethanol. Habitat descriptions and behavioral observations are based on fishes videotaped in situ and reported on by Tunnicliffe et al. (2010), with additional observations made from two fish appearing in a short video provided from the ROV ROPOS (Cruise Chief Scientist Dr. Ulrich Schwarz-Scham- 54 · Zootaxa 3061 © 2011 Magnolia Press MUNROE ET AL. pera) and edited by Jonathan Rose (University of Victoria; video available at http://ocean.si.edu/ocean-videos/ tonguefish-underwater-volcano-19). Size at maturity was determined from the stage of ovarian development in the female holotype following criteria outlined in Munroe (1998). Symphurus maculopinnis, new species Figures 1–7; Table 1 Symphurus species B. Tunnicliffe et al. 2010:4 (species recognition; bathymetric and ecological notes; genetic sequence data; distribution, abundance; color photographs). Holotype. USNM 398820, female, 84.4 mm SL, collected on Volcano–19 near Marker 40, Tonga Arc (24°48.439'S, 177°0.009'W), West Pacific Ocean, 561 m, collected with hand-held net (Sample Number R1048– 12) by ROV ROPOS directed by Anna Metaxas, 7 May 2007. FIGURE 2.Symphurus maculopinnis,n. sp. (Holotype USNM 398820; female, 84.4 mm SL), collected at 561 m on Volcano– 19, Tonga Arc, West Pacific. A. Ocular-side of alcohol-preserved specimen. B. Blind-side of same specimen. Diagnosis. Symphurus maculopinnis is distinguished from all other species of Symphurus by the combination of a 1–2–2–2–1 ID pattern, 14 caudal-fin rays, 3+6 abdominal vertebrae, 49 total vertebrae, about 89 scales in a longitudinal row, 92 dorsal-fin rays, 77 anal-fin rays, a blunt squarish snout, blind-side lower jaw with thick lip with conspicuous plicae, and by conspicuous ocellated (sometimes partially) black spots on the posterior dorsal and anal fins. Description. Morphological description based entirely on holotype (adult female, 84.4 mm SL; Figs. 1–5). A medium-sized species of Symphurus measuring to at least 100 mm SL [based on photographed specimens in Tun- nicliffe et al. (2010)]. ID pattern 1–2–2–2–1 (Figs. 3–4). Caudal-fin rays 14. Dorsal-fin rays 92. Anal-fin rays 76 (+1 fin ray missing). Pelvic-fin rays 4; posteriormost pelvic-ray connected to first anal-fin ray by delicate mem- NEW WESTERN PACIFIC TONGUEFISH Zootaxa 3061 © 2011 Magnolia Press · 55 brane. Total vertebrae 49; abdominal vertebrae 9 (3+6) (Fig. 4). Hypurals 5. Longitudinal scale rows about 89. Scale rows on head posterior to lower orbit 21. Transverse scale rows 45. Morphometrics (expressed as percentages of SL or HL; Table 1). Body moderately elongate (Fig. 2) with greatest body depth (29.6% SL) located in its anterior third (between anal-fin rays 10–25) and with gradual poste- rior taper. Preanal length slightly smaller than body depth. Head long, moderately wide, with squarish snout (Figs. 2A, 5); head length (21.6% SL) less than head width (25.6% SL; HW/HL = 1.20). Lower head lobe (12.7% SL) narrower than upper head lobe (16.2% SL); lower head lobe width less than postorbital length. Snout relatively broad with squarish anterior profile; snout length 18.1% HL; snout slightly longer than eye diameter. Dermal papil- lae conspicuously present on both ocular and blind sides of snout and head, but more numerous on blind-side head (Figs. 5A–B). Ocular-side snout and head with several short, vertical rows of dermal papillae as well as other TABLE 1. Morphometric features for holotype (USNM 398820) of Symphurus maculopinnisn. sp. and 19 S. ocellatus. SL in mm; characters 2–9 in % of SL; 10–17 in % of HL. S. maculopinnis S. ocellatus Character Holotype N=19 1. Standard length 84.4 75.6–120.5 2. Body depth 29.6 23.5–30.1 3. Preanal length 25.6 20.0–23.5 4. Head length 21.6 15.9–20.0 5. Head width 25.6 18.3–22.6 6. Postorbital length 13.5 10.5–13.7 7. Upper head lobe 16.2 9.9–15.6 8. Lower head lobe 12.7 7.4–11.9 9. Caudal-fin length 12.9 9.6–12.1 10. Postorbital length 62.6 63.0–68.5 11. Predorsal length 21.9 19.5–29.7 12. Snout length 18.1 15.6–21.7 13. Upper jaw length 25.8 20.3–24.8 14. Eye diameter 16.5 12.0–15.6 15. Chin depth 20.3 16.2–20.8 16. Upper opercular lobe 18.7 13.1–21.1 17. Lower opercular lobe 40.1 29.3–37.3 18. HW/HL 1.20 1.04–1.21 19. Pupil/Eye diameter 0.53 0.43–0.70 dermal papillae not arranged in obvious rows; ocular-side anterior head region dorsal to eyes with five, short, verti- cal rows of dermal papillae; anterior head region posterior to eyes with single, longer, horizontal row of dermal papillae; a single, short, vertical row of dermal papillae ventral to mid-region of lower eye; cheek region (ventral to lower jaw) with five, short, vertical rows of dermal papillae (Fig. 5A). Blind-side head and snout (Figs. 5B) with numerous, conspicuous, vertical and horizontal rows, sometimes interconnected, of dermal papillae extending nearly to ventro-posterior margin of opercle. Ocular-side anterior naris tubular, elongate; just reaching anterior margin of lower eye when depressed posteriorly. Posterior ocular-side naris a short, wide, tubular opening located nearly at vertical through anterior margin of pupil of lower eye. Blind-side anterior naris a short, slender, unpig- mented tube located anterior to vertical through mid-jaw and barely perceptible among dense patch of dermal papillae. Blind-side posterior naris a short, posteriorly-directed tube located at posterior margin of dense patch of dermal papillae dorsal to horizontal through base of anterior blind-side nostril. Posterior margin of ocular-side maxilla at vertical through anterior region of pupil of lower eye. Ocular-side lower jaw without fleshy ridge. Blind- side lower jaw with thick lip with numerous plicae. Chin depth (20.3% HL) slightly greater than snout length (18.1% HL). Lower eye large (16.5% HL); eyes slightly subequal in position, anterior margin of lower eye at point 56 · Zootaxa 3061 © 2011 Magnolia Press MUNROE ET AL. between verticals through anterior margin of upper eye and anterior margin of pupil of upper eye. Eyes separated by narrow interorbital space about 1–2 scales wide; dorsal aspects of eyes with 3–4 small, ctenoid scales; also with several rows of small ctenoid scales in interorbital space anteriorly. Pupillary operculum absent. Dorsal-fin origin at vertical through anterior margin of pupil of upper eye; predorsal distance 21.9% HL. Postorbital length 13.5% SL. Lower lobe of opercle (40.1% HL) twice as wide as upper opercular lobe (18.7% HL) and with its posterior margin extending noticeably further posteriorly than margin of upper opercular lobe. FIGURE 3. Radiograph of Symphurus maculopinnis,n. sp. (Holotype USNM 398820; female, 84.4 mm SL), collected at 561 m on Volcano–19, Tonga Arc, West Pacific. Both sides of dorsal- and anal-fin rays without scales. Distal tips of most dorsal- and anal-fin rays free from interradial membranes. Caudal fin short (12.9% SL), pointed, with middle fin rays slightly longer than others; basal halves on both sides of caudal fin with several rows of small, ctenoid scales. Ocular-side dentary and maxilla with few teeth on their anterior regions. Blind-side dentary with 2–3 rows of well-developed teeth anteriorly expanding to several rows posteriorly; blind-side premaxilla with 2 rows of well- developed teeth curving slightly inwards. Scales moderately large; ctenoid on both sides of body. Live color (Figs. 1; 6–7; based on the holotype and other specimens photographed or videographed in situ at ~433–561 m on Volcano–19 from the ROV ROPOS; video edited by Jonathan Rose). Ocular-side head and body grayish-green to light brown and overlain with numerous, irregular, dark-brown blotches of various sizes; also with several incomplete, wide (5–7 scale rows), darker crossbands in midbody region and one complete crossband on caudal 1/4th of body; also with several smaller (2–4 scales wide), irregular white markings scattered over body sur- face, the largest of which overlies the dorsoanterior abdomen; and with two darker patches– one dorsal to pelvic fin on dorsoanterior abdomen and another situated at ventroposterior margin of abdomen. Dorsal and anal fins each with a conspicuous, intense, ocellated (sometimes partially) black spot on their pos- teriormost rays (about the posteriormost 6–7 rays based on preserved specimen). Sometimes, black spots com- pletely ocellated by white ring; otherwise, black spot only partially surrounded by one or two much smaller, irregular, white spots situated around its outer periphery. White pigment of dorsal-fin ocellus frequently not com- pletely surrounding black spot near body nor as intense as ocellated ring and spots surrounding dark spot on anal fin. Dorsal and anal fins also with alternating series of lighter and darker blotches (about 4–6 fin-rays wide) throughout nearly entire length of fins. Dorsal-fin rays in anteriormost part of fin with distal thirds bright white and proximal two-thirds darker. Dorsal-fin rays in remainder of fin, and especially in more posterior blotches, more darkly pigmented over entire length of fin rays and also on fin membrane compared with those in blotches on ante- rior region of fin. Anal fin also with alternating series of lighter (some bright white) and darker blotches (about 4– NEW WESTERN PACIFIC TONGUEFISH Zootaxa 3061 © 2011 Magnolia Press · 57 6 rays in each series) throughout length of fin to just before black spot on posteriormost fin rays. Anal-fin rays throughout entire fin pigmented over most of their lengths. Anteriormost anal-fin rays nearly entirely white, con- trasting sharply to more heavily pigmented darker rays in remainder of fin. Anal-fin membrane in anterior region of fin pigmented only on its basal half, membrane in remainder of anal fin, especially that in dark blotches in posterior third of fin, more heavily pigmented nearly to its distal margin. Caudal fin dusky with several rays partially streaked with white and with several small whitish spots also on caudal-fin base. Pelvic-fin rays white throughout their lengths, in stark contrast to body color, especially contrasting against dark blotch on nearby abdomen. FIGURE 4. Radiograph of head and anterior body region of Holotype (USNM 398820) of Symphurus maculopinnis, n. sp., depicting 1–2–2–2–1 pattern of interdigitation of anterior dorsal-fin pterygiophores and neural spines and nine abdominal ver- tebrae. Note also the sediment in the alimentary tract visible in this xray. Color in alcohol (Figs. 2, 5; based only on holotype). Ocular-side background pigmentation uniformly medium dark brown with several diffuse, irregular, darker areas; region overlying abdomen dark, brownish-black (Fig. 2A). Ocular-side head and body scales with posterior halves of exposed region more darkly pigmented than anterior halves of exposed portions. Head coloration similar to that on body, except posterior margin of opercle out- lined in black (Fig. 5A). Snout nearly to distal tip with similar pigment as that on rest of head; distalmost snout 58 · Zootaxa 3061 © 2011 Magnolia Press MUNROE ET AL. region lighter than more posterior regions. Ocular-side lips dark black, except posteriormost parts of upper and lower lips lighter brown. Ocular-side anterior nostril black for most of its length; distalmost tip whitish. Ocular-side inner opercular lining white, ventral half with several faint melanophores; inner lining of blind-side opercle white. Isthmus on both sides with small melanophores, more concentrated on ocular-side isthmus than on blind-side coun- terpart. FIGURE 5. Close-up views of ocular (A)and blind (B) sides of head of Symphurus maculopinnis, n. sp. (Holotype USNM 398820; female, 84.4 mm SL) collected at 561 m on Volcano–19, Tonga Arc, West Pacific. Arrows indicate areas with conspic- uous rows of dermal papillae. Blind-side background coloration of head and body uniformly whitish except for dark, bluish-black abdomen (Figs. 2B, 5B). Dorsal and ventral margins of blind-side head (Fig. 5B) also with numerous, small, darkly-pig- mented melanophores on several rows of scales (similar in color to scales on ocular-side head); central portion of posterior region of blind-side head with irregular patch of numerous, faint, small melanophores (visible under mag- nification). Two nearly parallel, zig-zagging rows of clusters of faintly-pigmented melanophores extending posteri- orly from posterior margin of head nearly to caudal-fin base (Fig. 2B); dorsal row located about 1/4th body width from dorsal-fin base, ventralrow located medially just ventral to body mid-point. Single row of faint melanophores evident internally in dermis about at medial ends of proximal pterygiophores in both posterior 3/4ths of dorsal fin and for nearly entire length of anal fin. Ventroanterior region of blind-side abdomen dorsal to pelvic fin with dark patch of scales. Anal sphincter white. Blind-side body also with short, diffuse, continuous, dark sooty-brown smudge located dorsal to anal-fin rays 6–17 (corresponding to location of ovary). Ocular sides of dorsal and anal fins uniformly sooty-gray throughout most of their lengths. Most fin rays streaked with darker pigment than that on connecting membranes, distalmost tips of fin rays white (Fig. 2A). Pig- ment on rays and membranes progressively intensifying posteriorly in both fins. Dorsal and anal fins also with sin- gle, conspicuous, non-ocellated, black spot on their posteriormost rays. Anteriormost dorsal-fin rays streaked with dark pigment contrasting with lightly pigmented connecting membranes. Next several successive dorsal-fin rays also more darkly pigmented than connecting membranes, but membranes also feature clusters of melanophores in their distal sections. At about 1/3 length of fin and continuing posteriorly, nearly entire dorsal-fin membrane cov- NEW WESTERN PACIFIC TONGUEFISH Zootaxa 3061 © 2011 Magnolia Press · 59 ered with darker melanophores. Posteriormost six dorsal-fin rays with conspicuous black spot extending from fin base (and also slightly on body) to nearly distalmost tips of rays. Blind sides of anteriormost 3–4 dorsal-fin rays with speckling along entire lengths of rays and also on connect- ing membranes (Fig. 2B). Successive rays in anterior fin generally whitish with faint, small melanophores on rays and membrane. Pigmentation similar throughout remainder of fin, but becoming progressively darker posteriorly, and culminating with posteriormost rays bearing dark spot. Ocular sides of anteriormost anal-fin rays more lightly streaked than those of more posterior rays; their con- necting membranes only lightly pigmented in their distal regions. At about midpoint of anal fin, pigment covers entire interradial membrane. Seven posteriormost anal-fin rays covered by intense black spot extending from fin- ray bases nearly to distalmost tips. Blind side of anal fin generally whitish anteriorly, with rays in anterior one-third of fin with faint melano- phores; pigmentation progressively intensifying posteriorly, with darkest pigment on posteriormost fin rays bearing spot. Ocular sides of pelvic-fin rays with dark melanophores similar to those on anal fin; blind side of pelvic fin whitish, with faint, small, darker spots. Both sides of caudal fin uniformly sooty grayish-black throughout; ocular side of fin darker than blind side. Both sides of caudal fin with nearly entire lengths of rays and their connecting membranes heavily pigmented, and with distal thirds of central caudal-fin rays darker (nearly black) compared with their proximal two-thirds. Small, ctenoid scales on basal half of both sides of caudal fin with numerous, small, darkly-pigmented melanophores scattered over their exposed surfaces (best viewed under magnification). Distribution and habitat. Symphurus maculopinnis is known only from the holotype, collected on soft sedi- ments at 561 m (Fig. 1), and 56 other individuals observed, but not collected, on a variety of substrata, including coarse gravel, gravelly sand-shell and gravelly sand (Figs. 6–7), located between 433–561 m on Volcano-19, Tonga Arc, western Pacific. Tunnicliffe et al. (2010: Table 2) indicated that this species was also observed at shallower depths (between 195–381 m) at a second, nearby site (Volcano–1). However, re-examination of photographs indi- cates that the individuals upon which these observations were based can not be positively identified. Thus, reported occurrences of S. maculopinnis both at Volcano–1 and at these shallower depths is tenuous. Symphurus maculopinnis is considered to be a true vent species (Tunnicliffe et al. 2010) as it was most often observed (90% of 57 observations) within 30 m of point sources of obvious venting. It was observed to be abun- dant in areas near a high temperature vent that also supported abundant clams in the surrounding sediments. A probe inserted ca. 10 cm in the sediments registered 11°C (bottom water temperature 5°C), which indicated an area of low flux of hydrothermal fluid. Symphurus maculopinnis was not observed at other vent sites located deeper than 600 m either on Volcano–19 or at other volcanoes studied by Tunnicliffe et al. (2010). Etymology. The specific name, maculopinnis, from the Latin macula, meaning spot, and pinna, meaning fin, in reference to the conspicuous spots on the dorsal and anal fins of this species. To be treated as a noun in apposi- tion. Remarks. The holotype of S. maculopinnis, the only specimen collected and radiographed thus far (Figs. 3–4), features 14 caudal-fin rays, 9 abdominal vertebrae and a 1–2–2–2–1 ID pattern (Fig. 4). The 1–2–2–2–1 ID pattern is unusual among symphurine tonguefishes (Munroe 1992) as none of the 79+ nominal species of Symphurus examined to date feature a predominant 1–2–2–2–1 ID pattern (Munroe 1992; Munroe unpubl. data; Lee et al. 2009a; 2009b). Most species in the genus that have 14 caudal-fin rays and 9 abdominal vertebrae (N= 19 species) have either the 1–2–2–2–2 (14 species) or the 1–2–3–2–2 ID pattern (3 species) as their predominant pattern (Mun- roe 1992; Krabbenhoft & Munroe 2003; Munroe 2006; Munroe & Hashimoto 2008; Lee et al. 2009a, 2009b). Another seven species in the genus, also with 14 caudal-fin rays, have an ID pattern similar to that observed in the holotype of S. maculopinnis, where only a single pterygiophore inserts into interneural space 4 (1–2–2–1–2 ID pat- tern). However, four of these species also have 10 abdominal vertebrae (compared with only nine in S. maculopin- nis). Number of abdominal vertebrae, a very conservative character within Symphurus, is one of the most reliable meristic characters useful in diagnosing these species. Three other species, S. fuscus Brauer, S. macrophthalmus Norman and S. schultzi Chabanaud, have 9 abdominal vertebrae and 14 caudal-fin rays (Munroe 1992) and, based on the limited data available for these species, also appear to have an ID pattern featuring a 1–2–2–1–2 arrange- ment of pterygiophores, which is similar to that observed in S. maculopinnis. All three species have been rarely captured and the data are insufficient to determine whether this is their predominant ID pattern. The ID pattern of 60 · Zootaxa 3061 © 2011 Magnolia Press MUNROE ET AL. S. fuscus, for example,is known only from the holotype; that of S. macrophthalmus is known from the holotype and a paratype; and for S. schultzi, two patterns are represented among four paratypes (all with 1–2–2–1–2 ID pattern) and the holotype (with 1–2–2–2–2 ID pattern). FIGURE 6.In situ photographs of two individuals of Symphurus maculopinnis,n. sp., observed on Volcano–19, Tonga Arc, West Pacific, featuring different asymmetries. A. Specimen with sinistral asymmetry photographed at ca. 439 m. B. Frame grab of specimen featuring reversed (dextral) asymmetry videographed at ca. 560 m. Note also posture of both fish with raised cau- dal regions displaying prominent spots on posterior dorsal and anal fins as they swim over the bottom. Although the ID pattern of the holotype of S. maculopinnis features only a single pterygiophore inserted into the fifth interneural space, it is possible that this is not the predominant ID pattern for this species. More likely, the predominant ID formula for this species is 1–2–2–2–2. Among 14 species characterized by the 1–2–2–2–2 ID pat- tern, some individuals are observed, albeit infrequently, with a variant 1–2–2–2–1 ID pattern (6 of 258 individuals or 2.3% of specimens radiographed). Alternatively, if the predominant ID pattern for S. maculopinnis is found to be 1–2–2–2–1, then, based on what is presently known about ID patterns in symphurine tonguefishes, this species would feature a unique predominant ID pattern among species of Symphurus. Determination of the predominant ID pattern for S. maculopinnis, as for several other deepwater symphurine species, will have to await collection of additional specimens. Symphurus maculopinnis is the third tonguefish species known to inhabit hydrothermal vent areas, all of which are located on western Pacific Ocean volcanic arcs (Tunnicliffe et al. 2010). Other species found at hydrothermal vents include S. thermophilus (Munroe & Hashimoto, 2008), and undescribed “species A” recently discovered by Tunnicliffe et al. (2010). Munroeand Hashimoto (2008) described S. thermophilus, based on specimens collected at hydrothermal sites located in the North Pacific and South Pacific oceans. However, analysis of mitochondrial DNA samples and morphological comparisons (Tunnicliffe et al. 2010) reveal that two species are represented among specimens previously identified as S. thermophilus. Based on their results, Tunnicliffe et al. (2010) deter- NEW WESTERN PACIFIC TONGUEFISH Zootaxa 3061 © 2011 Magnolia Press · 61 mined that S. thermophilus is known at hydrothermal sites only in the northwest Pacific Ocean, while undescribed species A (including some specimens previously identified as S. thermophilus by Munroe & Hashimoto) occurs in the South Pacific Ocean on several volcanoes along the Kermadec–Tonga Arc between New Zealand and Samoa. FIGURE 7. In situ photographs of three individuals of Symphurus maculopinnis,n. sp. observed on Volcano–19, Tonga Arc, West Pacific, illustrating variation in live coloration, variety of substrata inhabited by this species, and interesting posture of raising the caudal region (indicated by arrows) thereby displaying prominent spots on the posterior dorsal and anal fins. A. Specimen with conspicuous, complete ocellus on dorsal and anal fins photographed on coarse pebbly substratum. B. Specimen (ca. 12.4 cm in length based on 10 cm distance between green laser dots) with incomplete white ring around black fin spots photographed on gravelly sand-shell substratum. C. Two specimens photographed on gravelly sand-shell substratum. Smaller specimen features complete ocellus on dorsal and anal fins and slight banding on body; larger specimen features little, if any, white pigment around periphery of black fin spots and distinct banding on body. At least 10 of the 79+ described species of Symphurus, including S. maculopinnis, have prominent (sometimes ocellated) spots on their dorsal, anal or caudal fins. These species live at a variety of depths ranging from 4 to 640 m (most occur in < 100 m) and inhabit environments from white, sandy substrata in shallow seagrass beds in clear tropical waters to various substrata in relatively dim lit waters on the upper continental slope and at seamounts. Of species with ocellated fins, only three occupy relatively deepwater habitats; S. stigmosus Munroe occurs from 192– 373 m (Munroe 1998), S. ocellatus occurs from 430-640 m (Munroe, unpub. data), and S. maculopinnis occurs between 433–561 m (Tunnicliffe et al. 2011). None of the deepest-dwelling (800–1500 m) species of Symphurus have ocellated fins. In a detailed study (1200 specimens, 50 species) of the occurrence of ocellated spots on dorsal fins in the deepsea ophidiid genus Neobythites Goode & Bean, Uiblein & Nielsen (2005) report 22 of 50 species in this genus with distinct ocelli and that these species occur at mean collecting depths ranging from 150 to 450 m, depths not all that different than those recorded for the three deepest-dwelling species of Symphurus with ocellated fins. Since a lower depth limit for color vision in clear oceanic waters is estimated to occur at about 550 m (Clarke & Denton 1961), the deeper depths where some of these species of Symphurus and Neobythites (Uiblein & Nielsen 62 · Zootaxa 3061 © 2011 Magnolia Press MUNROE ET AL.