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Accepted by M.T. Craig: 8 Aug. 2011; published: 18 Oct. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 3061: 53–66 (2011)
www.mapress.com/zootaxa/Article
53
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: munroet@si.edu
2Department of Biology, University of Victoria, PO Box 3080, Victoria, British Columbia, Canada V8W 3N5.
E-mail: verenat@uvic.ca; jentyler24@gmail.com
3Corresponding author. E-mail: munroet@si.edu
Abstract
Symphurus maculopinnis n. 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)
of S. 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).
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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
ROV ROPOS 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-
Zootaxa 3061 © 2011 Magnolia Press · 55
NEW WESTERN PACIFIC TONGUEFISH
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-
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56 · Zootaxa 3061 © 2011 Magnolia Press
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 maculopinnis n. sp. and 19 S. ocellatus. SL in
mm; characters 2–9 in % of SL; 10–17 in % of HL.
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
Character S. maculopinnis
Holotype S. ocellatus
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
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NEW WESTERN PACIFIC TONGUEFISH
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–
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58 · Zootaxa 3061 © 2011 Magnolia Press
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
Zootaxa 3061 © 2011 Magnolia Press · 59
NEW WESTERN PACIFIC TONGUEFISH
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, ventral row 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-
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60 · Zootaxa 3061 © 2011 Magnolia Press
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
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NEW WESTERN PACIFIC TONGUEFISH
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). Munroe and 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-
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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
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NEW WESTERN PACIFIC TONGUEFISH
2005) with ocellated fins occur may have natural light levels that are low to non-existent. Detection of visual sig-
nals, such as ocellated spots, in depths where light levels are low might appear to be limited in species inhabiting
such depths and their usefulness as signals questionable. However, Uiblein & Nielsen (2005) cited several studies
among shallow-water fishes living in reduced light conditions where visual acuity is low that provide evidence
these species may effectively use contrasting body coloration for signaling. Uiblein & Nielsen (2005) note further
that ocelli are among the most contrasting color patterns and hence should be visible in semidark habitats such as
those on the lower continental shelf and upper continental slope. This would apply equally well on Volcano–19
where the depths occupied by S. maculopinnis are in similar low-light environments.
No behavioral observations have been made on tonguefishes with ocellated fins regarding the function of such
conspicuous pigmentation features. For other fishes, ocellated pigment spots have been suggested to serve as spe-
cies recognition signals (Kohda & Watanabe 1990; Uiblein & Nielsen 2005) or as antipredator devices (Neudecker
1989: Hasson 1991; Uiblein & Nielsen 2005). During the course of exploration of hydrothermal vent areas and
other habitats on Volcano–19, 57 individuals of S. maculopinnis occurred in ROV imagery (Tunnicliffe et al.
2010). Examination of the behavior of these tonguefishes provides some insight into the function of ocellated fin
spots in this species. Tunnicliffe et al. (2010) note that individuals of S. maculopinnis appeared to be more quies-
cent than representatives of Symphurus species A observed in the same areas. Individual S. maculopinnis were
observed to flip sediment over their bodies to increase their camouflage, and when approached by another fish, one
S. maculopinnis raised its tail, thereby elevating the conspicuous eyespots on the dorsal and anal fins. We note two
other videographed individuals of S. maculopinnis closely associated with the coarse sedimentary substratum that
were initially at rest and partially buried (video available at http://ocean.si.edu/ocean-videos/tonguefish-underwa-
ter-volcano-19). They then began to swim slowly over the substratum propelled by flexing anterior to posterior
waves of muscle contraction down the body and dorsal and anal fins. While swimming, these two fish also exhib-
ited a similar posture of raised posterior body regions with dorsal- and anal-fin rays widely spread thereby conspic-
uously displaying the pigmented spots on these fins. Several other individuals observed and photographed on the
seamount also had raised caudal regions displaying their prominent ocellated fins (see Figs. 6A–B; 7B–C). Numer-
ous observations of individuals of S. maculopinnis displaying a swimming posture with eyespots prominently dis-
played when in the presence of conspecifics suggests that this posture serves as a species recognition signal among
conspecifics.
It is also possible that having eyespots so prominently displayed on up-raised and expanded fins also increases
the appearance of these tonguefishes beyond their actual size, whereby these ocellated spots would then function as
an antipredator mechanism. An enlarged appearance may deter visually hunting predators (Hasson 1991), or possi-
bly, as is the case with these tonguefishes with spots located on their posterior fins, the spots may confuse a preda-
tor into attacking the posterior end of the potential prey’s body where a vital zone is less likely to be struck
(Neudecker 1989; Hasson 1991), or where the attack zone may even allow the tonguefish to escape altogether as it
darts forward away from the predator’s point of attack at the rear of its body. Little is known about potential preda-
tors or predation rates on tonguefishes at hydrothermal vent areas (or elsewhere) and these aspects of their ecology
remain areas in need of further study.
Among videographed S. maculopinnis was a specimen featuring reversed (dextral) asymmetry (see Fig. 6B).
Based on observations from the short video of this specimen, it appears to be indistinguishable both in its morphol-
ogy, save for its reversed asymmetry, and its behavior from the other, non-reversed specimens of S. maculopinnis
appearing in this and other videos (Figs. 6A, 7A–C).
Capture of deepwater (>200 m) tonguefish with reversed asymmetry is rare, as only one other specimen, that of
the eastern Atlantic S. vanmelleae Chabanaud, is known (Munroe 1996). This video sequence of a reversed S. mac-
ulopinnis is also the first to observe behavior of a reversed individual of Symphurus, and represents the first
sequence to record in situ the reactions of conspecifics to behavior and movements of another conspecific with
reversed asymmetry.
Comparisons. Symphurus maculopinnis is only the second known Indo–West Pacific species of Symphurus
characterized by the combination of 14 caudal-fin rays, a 1–2–2–2–2 ID pattern (assuming this is predominant pat-
tern for the species; see Remarks Section above), and pigmented spots on its dorsal and anal fins. The only other
species of Symphurus featuring this combination of characters is S. ocellatus, known from deep waters (430–640
m) off East Africa (Heemstra 1986). Symphurus maculopinnis differs from S. ocellatus in having lower, and non-
overlapping, meristic features (total vertebrae 49 vs. 54–56, dorsal-fin rays 92 vs. 97–103, anal-fin rays 77 vs. 85–
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64 · Zootaxa 3061 © 2011 Magnolia Press
89), a blunt, squarish snout (vs. more pointed snout), thick blind-side lower lip with conspicuous plicae (vs. thin
blind-side lip without conspicuous plicae), longer (21.6% SL vs. 15.9–20.0% SL) and wider head (head width
25.6% SL vs. 18.3–22.6), and wider upper (16.2% SL vs. 9.9–15.6% SL) and lower (12.7% SL vs. 7.4–11.9% SL
in S. ocellatus) head lobes.
Other species of Symphurus with 14 caudal-fin rays and 50 or fewer total vertebrae include S. macrophthal-
mus, S. schultzi, S. monostigmus Munroe, S. bathyspilus Krabbenhoft and Munroe, S. multimaculatus Lee et al.,
and two other species that occur at hydrothermal vent areas, S. thermophilus and undescribed Species A, which is
closely related to S. thermophilus (Tunnicliffe et al. 2010). Symphurus maculopinnis differs from all of these other
species in having spots on the dorsal and anal fins. Symphurus maculopinnis differs further from S. macrophthal-
mus, a rarely-captured species known only from the holotype and a paratype (in pieces) collected between 457–549
m in the Gulf of Aden (Norman 1939), in having a smaller eye diameter (ED) (ED 16.5% HL vs. 21.1% HL in S.
macrophthalmus).
Symphurus maculopinnis differs further from S. monostigmus, known from two specimens collected in rela-
tively shallow waters (65–100 m) off East Africa (Munroe 2006), in having a darkly-pigmented ocular side with
conspicuous dark blotches and incomplete crossbands (vs. ocular side lightly pigmented with numerous freckles in
S. monostigmus). Symphurus maculopinnis also has a larger pupil (pupil diameter/eye diameter = 53% compared
with that of S. monostigmus (28–32%), a longer, narrower head (HW/HL =1.20 vs. 1.39–1.42), a shorter postorbital
length (POL) relative to head length (POL/HL =62.6% vs. 75.5–80.5%), a longer snout (18.1% HL vs. 14.5–17.2%
HL) and larger eye (ED = 16.5% HL vs. 10.0–13.8% HL), several small scales in the interorbital space and on
upper aspects of the eyes (scales absent in these areas in S. monostigmus), and S. maculopinnis has a wider lower
opercular lobe compared with the upper opercular lobe (vs. upper opercular lobe wider than lower opercular lobe in
S. monostigmus).
Symphurus maculopinnis differs from S. bathyspilus, a deepwater (248–500 m) species occurring in the Philip-
pine Archipelago and Indonesia (Krabbenhoft & Munroe 2003), in its squarish anterior head profile (vs. longer,
pointed snout profile) and different blind-side pigmentation (white with relatively few melanophores arranged in
zig-zag longitudinal rows vs. blind-side white with numerous melanophores scattered over blind-side surface and
heavier covering of melanophores primarily over pterygiophore regions of dorsal and anal fins). Symphurus macu-
lopinnis may have fewer dorsal- and anal-fin rays than does S. bathyspilus, as counts for these features in the holo-
type only overlap the lower end of ranges of these meristic features in S. bathyspilus. Additionally, the holotype of
S. maculopinnis has 49 total vertebrae and 45 scales in a transverse row versus the 50–54, usually 51–52, total ver-
tebrae and 29–34 scales in a transverse row in S. bathyspilus. Several morphometrics of the holotype also differ
from those of S. bathyspilus including its deeper body (body depth 29.6% SL vs. 21.8–25.8% SL), a smaller POL
(62.6% HL vs. 63.6–71.6% HL), and a wider upper head lobe (16.2% HL vs. 8.9–13.0% HL). The dorsal-fin origin
(anterior margin of pupil of upper eye) is located more anteriorly on the holotype of S. maculopinnis than is that of
S. bathyspilus (located between verticals through middle and posterior margin of upper eye).
Most meristic and morphometric features of S. maculopinnis overlap those of S. schultzi, a deepwater species
collected between 479–1013 m in the Philippine Archipelago (Chabanaud 1955), which is known only from four
paratypes (all with 1–2–2–1–2 ID pattern) and the holotype (with 1–2–2–2–2 pattern). The holotype of S. macu-
lopinnis differs in having 89 scales in a longitudinal series compared with 70–80 longitudinal scales in S. schultzi.
The two species also differ in the anterior profile of the head (squarish in S. maculopinnis vs. more pointed in S.
schultzi). The color patterns of the two species are also different. Chabanaud (1955) indicated the type series of S.
schultzi featured an evenly pigmented reddish-brown ocular side (specimens now faded), which is distinctly differ-
ent from that observed in the holotype and other specimens videographed of S. maculopinnis.
Symphurus maculopinnis has similar meristic features to those of S. multimaculatus Lee et al. collected in deep
waters (exact depths unknown, but thought to occur deeper than 300 m) off Taiwan (Lee et al. 2009a), but differs
distinctly from this species in its squarish anterior profile (vs. pointed anterior profile) and more elongate body
with greatest depth occurring over larger part of mid-body with gradual anterior and posterior taper (vs. greatest
body depth occurring over shorter part of mid-body with rapid anterior taper in S. multimaculatus). The two species
also differ in aspects of their blind-side coloration including the white opercle with few melanophores in S. macu-
lopinnis (vs. bluish-black blind-side opercle in S. multimaculatus) and the blind-side body mostly white with zig-
zag arrangement of melanophores (vs. blind side with dense concentrations of blackish-brown melanophores pre-
dominately covering pterygiophore regions of dorsal and anal fins). Symphurus maculopinnis has fewer longitudi-
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NEW WESTERN PACIFIC TONGUEFISH
nal and transverse scale rows (89 vs. 102–108, and 45 vs. 50–51) than does S. multimaculatus. Differences in the
morphology of these species are also evident in: ratios of head width to head length (HW/HL 1.20 vs. 1.26–1.50,
reflecting that S. maculopinnis has a shorter and narrower head); in eye diameters (16.5% HL in S. maculopinnis
vs. 9.1–10.1% HL); postorbital head lengths (shorter in S. maculopinnis 62.6% HL vs. 67.8–71.7% HL in S. multi-
maculatus); and pupil size (pupil diameter 53% of eye diameter in S. maculopinnis vs. 61–72% in S. multimacula-
tus).The two other tonguefishes captured at western Pacific hydrothermal vents, S. thermophilus (Munroe &
Hashimoto 2008) and the closely-related and undescribed Symphurus Species A of Tunnicliffe et al. (2010) from
the Kermadec Ridge, also have 14 caudal-fin rays, similar ID patterns, and overlapping counts for total vertebrae
and dorsal- and anal-fin rays with those observed in S. maculopinnis. Neither species has spots on the dorsal and
anal fins so they can readily be distinguished from S. maculopinnis as long as pigment is present on fins of the
specimens. Symphurus maculopinnis differs further from both species in its more elongate body with gradual taper
(vs. greatest body depth in anterior third of body with more rapid posterior taper), in having larger scales (89 in lon-
gitudinal series vs. 100–112), a larger lower opercular lobe (40.1% HL vs. 26.9–36.4% HL), smaller upper opercu-
lar lobe (18.7% HL vs. 21.7–36.4% HL), and shorter predorsal length (21.9% HL vs. 24.2–31.9% HL in these
others). Additionally, genetic data also support the hypothesis that S. maculopinnis differs distinctly from these
other species as Tunnicliffe et al. (2010) found a genetic distance of 30% (COI) and 14% (16s rRNA) between S.
maculopinnis and S. thermophilus/Symphurus species A.
Acknowledgments
L. Willis provided radiographs of the holotype and L. Willis and M. Nizinski provided radiographs of specimens
used in the comparisons. J. Clayton and D. Smith assisted with accessioning and cataloguing the holotype. J. Rose
compiled the videotape of specimens filmed in situ at Volcano–19 and provided field data associated with the holo-
type; C. Marzec and M. Benson assisted with publishing and archiving the tonguefish video to the Smithsonian
Institution’s Ocean Portal website; C. Ames assisted with making video grabs from video tapes. S. Raredon pro-
vided photographs of the holotype. Appreciation is extended to curators and collection staff who provided assis-
tance locating specimens, providing catalogue information, or who arranged for loan of specimens used in this
study. We acknowledge the field team who collected and filmed this species during the SO192–2 MANGO Expedi-
tion, particularly S. K. Juniper and C. Stevens; expedition co-chiefs were U. Schwarz-Schampera and M. Hanning-
ton with R/V Sonne and ROV ROPOS. Collection of this specimen was made possible by funding provided to V.
Tunnicliffe by NSERC Canada.
Literature cited
Chabanaud, P. (1955) Flatfishes of the genus Symphurus from the U.S.S. “Albatross” Expedition to the Philippines, 1907–
1910, Journal of the Washington Academy of Sciences, 45(1), 30–32.
Clarke, G.L, & Denton, E.J. (1961) Light and Animal Life. In: Hill, M.N. (ed.), The Sea. Wiley, New York, pp. 456–468.
Hasson, O. (1991) Pursuit-deterrent signals: communication between prey and predator. Trends in Ecology and Evolution, 6,
325–329.
Heemstra, P.C. (1986) Family Cynoglossidae. In: Smith, M.M., & Heemstra, P.C. (Eds.), Smith’s Sea Fishes, 1st ed. Southern
Book Publishers, Johannesburg, pp. 865–868.
Kohda, Y., & Watanabe, M. (1990) The aggression-releasing effect of the eye-like spot of the oyanirami Coreoperca kawame-
bari, a freshwater serranid fish, Ethology, 84, 162–166.
Krabbenhoft, T.J., & Munroe, T.A. (2003) Symphurus bathyspilus: a new cynoglossid flatfish (Pleuronectiformes: Cynoglossi-
dae) from deepwaters of the Indo–West Pacific, Copeia, 2003(4), 810–817.
Lee, M.-Y., Munroe, T.A., & Chen, H.-M. (2009a) A new species of tonguefish (Pleuronectiformes: Cynoglossidae) from Tai-
wanese waters, Zootaxa, 2203, 49–58.
Lee, M.-Y., Shao, K.-T., & Chen, H.-M. (2009b) A new species of deep-water tonguefish Genus Symphurus (Pleuronecti-
formes: Cynoglossidae) from Taiwan, Copeia, 2009(2), 342–347.
Munroe, T.A. (1992) Interdigitation pattern of dorsal-fin pterygiophores and neural spines, an important diagnostic character
for symphurine tonguefishes (Symphurus: Cynoglossidae: Pleuronectiformes), Bulletin of Marine Science, 50(3), 357–
403.
MUNROE ET AL.
66 · Zootaxa 3061 © 2011 Magnolia Press
Munroe, T.A. (1996) First record of reversal in Symphurus vanmelleae (Pleuronectiformes: Cynoglossidae), a deep-water
tonguefish from the tropical eastern Atlantic, Cybium, 20(1), 47–53.
Munroe, T.A. (1998) Systematics and ecology of tonguefishes of the genus Symphurus (Cynoglossidae: Pleuronectiformes)
from the western Atlantic Ocean, Fishery Bulletin, 96(1), 1–182.
Munroe, T.A. (2006) New western Indian Ocean tonguefish (Pleuronectiformes: Cynoglossidae, Symphurus), Copeia, 2006(2),
230–234.
Munroe, T.A., & Hashimoto, J. (2008) A new western Pacific tonguefish (Pleuronectiformes: Cynoglossidae): The first pleu-
ronectiform discovered at active hydrothermal vents, Zootaxa, 1839, 43–59.
Neudecker, S. (1989) Eye camouflage and false eyespots: chaetodontid responses to predators, Environmental Biology of
Fishes, 25, 143–157.
Norman, J.R. (1939) Fishes. The John Murray Expedition. 1933–34. Scientific Reports of the John Murray Expedition, 7, 1–
116.
Stoffers, P., Worthington, T.J, Schwarz-Schampera, U., Hannington, M.D., Massoth, G.J., Hekinian, R., Schmidt, M., Lundsten,
L.J., Evans, L.J., Vaiomo’unga, R., & Kerby, T. (2006) Submarine volcanoes and high-temperature hydrothermal venting
on the Tonga arc, southwest Pacific, Geology, 34, 453–456.
Tunnicliffe, V., Koop, B.F., Tyler, J., & So, S. (2010) Flatfish at seamount hydrothermal vents show strong genetic divergence
between volcanic arcs, Marine Ecology, 31(Suppl. 1), 1–9.
Uiblein, F., & Nielsen, J.G. (2005) Ocellus variation and possible functions in the genus Neobythites (Teleostei: Ophidiidae),
Ichthyological Research, 52, 364–372.
