Content uploaded by Mao-Ying Lee
Author content
All content in this area was uploaded by Mao-Ying Lee on Dec 12, 2015
Content may be subject to copyright.
A New Species of Deep-Water Tonguefish Genus Symphurus
(Pleuronectiformes: Cynoglossidae) from Taiwan
Mao-Ying Lee
1,2
, Hong-Ming Chen
2
, and Kwang-Tsao Shao
1
A new symphurine tonguefish species, Symphurus megasomus, is described on the basis of 18 specimens collected in deep
waters (471–640 m) off eastern Taiwan. This species is characterized by the combination of a predominant 1-2-3-2-2
pattern of interdigitation of dorsal pterygiophores and neural spines, 14 caudal-fin rays, 106–111 dorsal-fin rays, 91–96
anal-fin rays, 55–58 total vertebrae, 5 hypurals, deep body (27.2–30.8% SL), small (5.3–7.9% HL), round and separate
eyes, bluish-black opercle and peritoneum on both sides of body (in both fresh and alcohol-preserved specimens),
uniformly straw-colored to dark-brown ocular-side background pigmentation without freckles or mottled pigmenta-
tion, and uniformly white to light yellow blind side. Among congeners, S. megasomus is most similar to S. undatus, which
has the same ID pattern and number of caudal-fin rays, but differs markedly from that species in having more dorsal-
and anal-fin rays and in its ocular-side coloration (uniformly straw-colored to dark brown vs. freckles on the ocular side
of S. undatus).
T
HREE species of symphurine tonguefishes have been
reported from relatively deep waters (200–600 m)
around Taiwan (Shen, 1983, 1993; Shen and Lin,
1984). Two of these species (Symphurus orientalis, S. novem-
fasciatus) feature 12 caudal-fin rays, while the third species
(S. strictus) possesses 14 caudal-fin rays. Symphurus strictus is a
widespread species that is distributed from off the east coast
of Africa, (Heemstra, 1986 ), Japan (Och iai, 1984), and
Hawaii (Gilbert, 1905). Since the original description of S.
strictus by Gilbert (1905), many Indo-Pacific specimens of
several species of Symphurus with 14 caudal-fin rays have
been identi fied as this species (Munroe, pers. comm.).
However, meristic features for specimens identified as S.
strictus (Munroe, 1992) have the largest range among Indo-
Pacific Symphurus, thus it is difficult to confirm the accuracy
of identifications based solely on literature sources for
specimens previously identified as this species.
Munroe (1992) considered that, in addition to the number
of caudal-fin rays, the number of proximal dorsal-fin
pterygiophores inserting into each of the anterior five
interneural spaces (ID pattern) is a reliable diagnostic
character for identifying species of Symphurus. In many
cases, this character has been used to diagnose phenetically
similar species that overlap in fin-ray counts, but that have
different ID patterns. Based on this, we began checking the
ID patterns of Taiwanese specimens of Symphurus to assist
with identifications. After examination of all specimens
collected by the R/V Ocean Researcher I during a six-year
project (August 2001–July 2007) and others caught by deep-
sea commercial trawlers, we found 18 specimens that we
tentatively identified as S. strictus. Further investigation
revealed that these specimens had a different ID pattern, fin-
ray counts, and coloration compared with those of S. strictus.
These specimens, instead, were more similar in ID pattern
and fin-ray counts to two other western Pacific species, S.
undatus and S. hondoensis (Gilbert, 1905; Chabanaud, 1956;
Munroe, 199 2; Munroe and Amaoka, 1998). However,
differences in ocular-side pigmentation, together with
meristic and morphometric characters revealed that these
specimens represented an undescribed species.
MATERIALS AND METHODS
Some of the Symphurus specimens were collected by the R/V
Ocean Researcher I during a six-year project (August 2001–
July 2007) sponsored by the National Science Council;
others were caught by deep-sea commercial trawlers. A total
of 18 specimens of S. megasomus were radiographed in this
study. Methods for meristic and morphometric characters
and general terminology follow Munroe (1998). Terminol-
ogy for interdigitation patterns of proximal dorsal pterygio-
phores and interneural spines (ID pattern) follow Munroe
(1992). Morphometric characters were taken to the nearest
0.1 mm with dial calipers or ocular micrometer. Morpho-
metric features are expressed either as measurements in
percent standard length (SL) or percent head length (HL).
Coloration and pigmentation descriptions are based on
fresh specimens and those fixed in 10% neutralized formalin
and preserved in 70% alcohol. Maturity was estimated by
macroscopic examination of the extent of posterior elonga-
tion of ovaries and presence of developing ova in the ovaries
(both observed by using light transmitted through the
body). In species of Symphurus, no obvious differences were
apparent in testes size between mature and immature males;
therefore, estimates of maturity were based entirely on
females. Institutional abbreviations follow http://www.asih.
org/codons.pdf, with ASIZP (formerly ASIZT) for Biodiversity
Research Center, Academia Sinica, Taipei, Taiwan. Compar-
ative materials for all other Indo-Pacific species of Symphurus
were listed in Shen (1993), Munroe (1992), Munroe and
Amaoka (1998), Krabbenhoft and Munroe (2003), Munroe
(2006), and Munroe and Hashimoto (2008).
Symphurus megasomus, new species
Giant Tonguefish
Figures 1, 2; Tables 1, 2
Copeia cope-09-02-18.3d 3/3/09 03:55:59 342 Cust # CI-08-080R1
1
Laboratory of Fish Ecology and Evolution, Biodiversity Research Center, Academia Sinica, Nankang, 11529 Taipei, Taiwan; E-mail: (M-YL)
coleopetera@gmail.com; and (K-TS) zoskt@gate.sinica.edu.tw. Send reprint requests to K-TS.
2
Department of Aquaculture, National Taiwan Ocean University, No. 2, Peining Rd., 20224 Keelung, Taiwan; E-mail: (H-MC)
hmchen@mail.ntou.edu.tw.
Submitted: 2 May 2008. Accepted: 1 December 2008. Associate Editor: C. J. Ferraris.
F 2009 by the American Society of Ichthyologists and Herpetologists
DOI: 10.1643/CI-08-080
Copeia 2009, No. 2, 342–347
Symphurus strictus (not of Gilbert).—Shen, 1983:108, fig. 3;
Shen, 1984:141, pl. 141, fig. 442–10; Shen, 1993:581, pl.
197, fig. 3.
Holotype.—ASIZP 67640, male, 136.2 mm SL, off northeast-
ern coast of Taiwan, bottom trawl fishing, Da-shi Fish
Market, 5 February 2007, M.-Y. Lee.
Paratypes.—17 specimens, 88.2–146.7 mm SL: ASIZP 63163,
2, males, 121.0–136.2 mm SL, off Su-ao, eastern Taiwan,
beam trawl, 520–640 m, 24u51.799–24u49.839N, 122u02.549–
122u02.399E, Ocean Researcher I, CP 120, 31 July 2001. ASIZP
63845, 2, male and mature female, 110.5 and 129.6 mm SL,
respectively, off Su-ao, eastern Taiwan, beam trawl, 570 m,
24u52.159–24u49.639N, 122u03.129–122u02.669E, Ocean Re-
searcher I, CP 195, 11 September 2002. ASIZP 63160, 2,
mature females, 102 .2–121.9 mm SL, off Su-ao, eastern
Taiwan, otter trawl, 471–531 m, 24u48.479–24u51.929N,
122u02.409–122u02.449E, Ocean Researcher I, CP 120, 1
August 2001. MNHN 2008-1930 (formerly ASIZP 67636),
female, 127.0 mm SL, off northeastern coast of Taiwan,
bottom trawl, Da-shi Fish Market, 18 July 2006, M.-Y. Lee.
BSKU 96068 (formerly ASIZP 67637), male, 139.2 mm SL, off
northeastern coast of Taiwan, bottom trawl, Da-shi Fish
Market, 21 August 2006, M.-Y. Lee. BSKU 96069 (formerly
ASIZP 67638), male, 118.0 mm SL, off northeastern coast of
Taiwan, bottom trawl, Da-shi Fish Market, 21 August 2006,
M.-Y. Lee. MNHN 2008-1929 (formerly ASIZP 67643), male,
118.6 mm SL, off northeastern coast of Taiwan, bottom
trawl, Da-shi Fish Market, 9 March 2007, M.-Y. Lee. USNM
393588 (formerly ASIZP 67644), 4, 2 males and 2 mature
females, 123.3–146.7 mm SL, o ff northeastern coast of
Taiwan, bottom trawl, Da-shi Fish Market, 16 March 2007,
M.-Y. Lee. AMS I.44690-001 (formerly ASIZP 67649), 3, 2
males, 90.9 and 96.8 mm SL, respectively, and 1 immature
female, 88.2 mm SL, off northeastern coast of Taiwan,
bottom trawl, Da-shi Fish Market, 29 June 2007, M.-Y. Lee.
Diagnosis.—Symphurus megasomus is distinguished from all
congeners, except S. hondoensis and S. undatus,byits
predominant 1-2-3-2-2 ID pattern (vs. ID patterns different
in these other species). Symphurus megasomus differs from
both S. hondoensis and S. undatus in its smaller eye (5.3–7.9%
HL vs. 11.0–14.9% HL in
S. hondoensis and 11.5–13.8% HL in
S. undatus). Symphurus megasomus further differs from S.
hondoensis in having nine abdominal vertebrae (vs. 10 in S.
Copeia cope-09-02-18.3d 3/3/09 03:56:00 343 Cust # CI-08-080R1
Fig. 1. Symphurus megasomus, holotype, ASIZP 67640. (A) Ocular side. (B) Blind side.
Lee et al.—New species of Symphurus 343
hondoensis). From S. undatus, S. megasomus is also distin-
guished by its higher fin-ray counts (106–111 dorsal-fin rays
and 91–96 anal-fin rays vs. 101–105 dorsal-fin rays and 87–
91 anal-fin rays in S. undatus) and uniform straw-colored to
dark brown ocular-side coloration (vs. freckled ocular-side
pigmentation in S. undatus).
Description.—Symphurus megasomus is a large-sized species,
reaching sizes extending to 146.7 mm SL. Meristic charac-
ters are summarized in Table 1. Predominant ID pattern 1-2-
3-2-2 (12/18 specimens 5 67%). Caudal-fin rays 14 (only
one specimen had 12). Dorsal-fin rays 106–111, usually 108–
109. Anal-fin rays 91–96, usually 93–94. Pelvic-fin rays 4.
Total vertebrae 55–58 ; abdominal vertebrae 9 (3 + 6).
Hypurals 5. Longitudinal scale rows 108–119. Scale rows
on head posterior to lower orbit 20–22. Lateral scale rows
48–55, usually 49–51.
Proportions of morphometric features are presented in
Table 2. Body deep; maximum depth in anterior one-third
of body; body depth tapering gradually from anterior to
posterior. Preanal length smaller than body depth. Head
wide; head width greater than head length (HW/HL 1.38–
1.76, mean 5 1.54). Upper head lobe wider than lower head
lobe and longer than postorbital length. Lower opercular
lobe of ocular side wider than upper opercular lobe. Snout
round to nearly square anteriorly, its length greater than eye
diameter (SNL/ED 1.71–3.51, mean 5 2.73). Dermal papillae
large and present on blind-side snout and chin. Ocular-side
anterior nostril tubular and short (damaged in most
specimens), usually not reaching anterior margin of lower
eye when depressed posteriorly. Ocular-side posterior nostril
a small rounded tube located between upper and lower eye.
Blind- side anterior nostril tubular and elongate, easily
distinguishable from dermal papillae; blind-side posterior
nostril a shorter posteriorly directed tube situated posterior
to vertical at rear margin of jaws. Jaws short; posterior
margin of upper jaw usually reaching point between
verticals through anterior margin and midpoint of lower
eye. Eyes separate and small; with two to four rows of small
ctenoid scales in interorbital space, and with some small
scales covering upper aspects of eye. Pupillary operculum
absent. Dorsal-fin origin located anterior to vertical through
anterior margin of upper eye; second dorsal-fin ray located
at vertical through anterior margin of upper eye. No scales
present on dorsal- and anal-fin rays on either side. Caudal
fin short, without ctenoid scales on base of fin.
Three to four rows of well-developed teeth on blind-side
dentary; three rows of well-developed teeth on blind-side
premaxilla. One row of scattered teeth anteriorly on ocular-
side dentary and premaxilla; teeth absent on posterior
ocular-side dentary and premaxilla.
Coloration and pigmentation.—Ocular-side background gen-
erally straw-colored to dark-brown. Some specimens with
irregular, darkly shaded areas on ocular side. Head colora-
tion similar to that on body. Outer surface of both opercles
bluish-black. Posterior margin of opercle darker brown and
easily distinguishable from blue regions of opercle. Inner
surfaces of opercles with small dark brown or black dots. Lips
Copeia cope-09-02-18.3d 3/3/09 03:56:36 344 Cust # CI-08-080R1
Fig. 2. Radiographs depicting interdigitation patterns (ID patterns) of
proximal dorsal-fin pterygiophores and neural spines and numbers of
abdominal vertebrae for three species of Symphurus. (A) Symphurus
megasomus (ASIZP 67640) with 1-2-3 ID pattern and 9 (3+6)
abdominal vertebrae; (B) Symphurus hondoensis (ASIZP 61782) with
1-2-3 ID pattern and 10 (3+7) abdominal vertebrae; and (C)
r
Symphurus strictus (ASIZP 67639) with 1-2-2 ID pattern and 9 (3+6)
abdominal vertebrae.
344 Copeia 2009, No. 2
and chin region light brown, margins of lips pigmented by
small black dots. Ocular-side anterior nostril yellowish-brown.
Upper aspects of eyes, pupil, and eye socket bluish. Blind side
generally white to light yellow. Some specimens with
irregular, darker regions distributed over body, but pigment
patches not forming distinct blotches. Peritoneum bluish-
black, clearly observed through abdominal wall on both sides.
Colorat ion of alcohol-preserved specimens similar to
freshly caught fishes except that coloration of opercle and
peritoneal regions (on both sides) of the freshly caught
specimens with bluish-black background coloration blend-
ing more gradually into the brown color of the background.
Fin rays of dorsal, anal, and pelvic fins uniformly light
brown with dark brown or black margins; basal regions of fin
rays bluish. Numerous small spots covering entire fin
membranes on both sides of fin, which give the fin
membranes a bluish color. Coloration of blind-side fin rays
white and bluish along their basal margins. Alcohol-
preserved specimens with bluish color of fin membranes
faded and becoming light brown.
Size and sexual maturity.— Specimens examined were 88.2–
146.7 mm SL. One specimen is an immature female
(88.2 mm SL) that shows little elongation of the ovaries.
Seven specimens were mature females with elongate ovaries,
five (102.2–139.2 mm SL) were not gravid, while two
(129.6–143.4 mm SL) were gravid with obvious ova. Males
(n 5 10, 90.9–146.7 mm SL) attain similar size to those of
females.
Remarks.—Twelve of 18 specimens have a 1-2-3-2-2 ID
pattern. Another three specimens have a 1-2-3-3-2 ID
pattern, while one specimen each has a 1-2-2-3-2, 1-2-4-2-
2, and 1-3-3-2-2 ID pattern. All of these other patterns,
considered to be variants of the 1-2-3-2-2 ID pattern
(Munroe, 1992), are not found in species featuring other
ID patterns.
Distribution and ecology.—Symphurus megasomus is currently
known only from the continental shelf and upper conti-
nental slope off northeastern and eastern Taiwan. Twelve
specimens were collected by deep-sea trawlers operating in
the commercial fishery in Dashi, northeastern Taiwan;
research vessels collected six other specimens in this general
area. This species has not been found in the Donggang
fishport, which has landings from deep-sea commercial
trawlers fishing in the waters off southern Taiwan. Also, no
specimens of S. megasomus were collected off southwestern
Copeia cope-09-02-18.3d 3/3/09 03:56:41 345 Cust # CI-08-080R1
Table 1. Frequency of Meristic Characters of Symphurus megasomus. Counts for the holotype (ASIZP 67640) indicated by an asterisk (*).
ID Pattern
1-2-3-2-2* 1-2-2-3-2 1-2-3-3-2 1-3-3-2-2 1-2-4-2-2 n
12 1 3 1 1 18
Caudal-fin rays
12 13 14* n
1 0 17 18
Dorsal-fin rays
106 107 108 109* 110 111 n
12940218
Anal-fin rays
91 92 93 94* 95 96 n
12 7 6 1 118
Total vertebrae
55 56 57* 58 n
17 9 118
Longitudinal scale count
108 109 110* 111 112 113 114 115 116 117 118 119 n
101 21210133116
Head scale count
20* 21 22 n
76215
Lateral scale count
48 49* 50 51 52 53 54 55 n
1 4 2 4011114
Lee et al.—New species of Symphurus 345
Taiwan by research vessels involved in the deep-sea
investigation conducted in this region. Symphurus mega-
somus is a deep-water tonguefish with specimens collected
from 471–640 m by research vessels. Specimens were found
in the landings at Dashi fishport, which also contained S.
orientalis, S. strictus, and other deep-sea fishes that live at
about 200–400 m. The community of fishes comprising the
landings at the Dashi fish port suggests that the depth range
of S. megasomus may be greater than that indicated by the
research vessel collections.
Polychaetes, small crustaceans and some unidentified
detritus were found in the digestive systems of fishes
examined. Little else is known regarding the biology of this
species.
Etymology.—The name megasomus is derived from the Greek,
mega, meaning large, and somus, meaning body, in reference
to the large size of this species and the deep body without
obvious tapering in the posterior body.
DISCUSSION
Among Indo-Pacific symphurine tonguefishes, 20 described
species possess 14 caudal-fin rays (Munroe, 1992; Krabben-
hoft and Munroe, 2003; Munroe, 2006; Munroe and
Hashimoto, 2008). Of these 20 species, 17 are readily
distinguished in having different predominant ID patterns.
Only two species, S. hondoensis and S. undatus, share a 1-2-3-
2-2 ID pattern (Fig. 2A) with S. megasomus.
Among species with a predominant 1-2-3-2-2 ID pattern,
S. megasomus has nine (3+6) abdominal vertebrae compared
with that of S. hondoensis, which has ten (3+7) abdominal
vertebrae (Fig. 2B). Symphurus megasomus also has a much
smaller eye (5.3–7.9% HL vs. 11.0–14.9% HL) and a much
deeper body (27.2–30.8% SL vs. 21.4–26.8% SL) compared
with those of S. hondoensis (Lee, unpubl. data). Symphurus
megasomus can also be distinguished from S. hondoensis in its
white to light yellow blind side (vs. blind side with dark
yellowish-green to dark brown pigmentation in S. hondoen-
sis) and absence of pigmentation on the body regions
overlying the dorsal and anal pterygiophores (vs. pigment
present in this region in S. hondoensis).
Among other Indo-Pacific Symphurus, S. megasomus is also
similar in ID pattern to S. undatus, a deep-water tonguefish
known fro m the Philippines and Haw aii (Munroe and
Amaoka, 1998). Symphurus megasomus generally has smaller
eye (11.5–13.8% HL in S. undatus) and higher fin-ray counts
than does S. undatus (106–111 dorsal-fin rays, 91–96 anal-fin
rays vs. 101–105 dorsal-fin rays, 87–91 anal-fin rays). These
two species are readily distinguished by differences in their
ocular-side pigmentation. Symphurus m egasomus has a
uniformly straw-colored to dark brown ocular side, whereas
in S. undatus the ocular side is freckled or mottled.
We also compared features of S. megasomus with other
Indo-Pacific species that have similar fin-ray counts but
different predominant ID patterns. Symphurus megasomus
overlaps in vertebrae and fin-ray counts with S. strictus. The
ID pattern (Fig. 2C) usually distinguished these two species
(1-2-3-2-2 in S. megasomus vs. 1-2-2-2-2 in S. strictus). In the
Taiwan region, counts for S. megasomus do not overlap with
those of S. strictus (113–116 dorsal-fin rays, 98–104 anal-fin
rays, 60–62 vertebrae); nevertheless, these species still have
been misidentified in past studies (Shen, 1983, 1984, 1993).
In S. megasomus, the opercle is bluish-black rather than the
brown opercle of S. strictus. The range in fin-ray counts and
geographic distribution of S. strictus is large (Munroe, 1992),
and it is possible that a species complex is present among
populations now identified as S. strictus.
Copeia cope-09-02-18.3d 3/3/09 03:56:42 346 Cust # CI-08-080R1
Table 2. Morphometrics for Holotype (ASIZP 67640) and 17 Paratypes of Symphurus megasomus. SL in mm; characters 2–15 in percent SL; 16–22
in percent HL; n 5 number of specimens measured.
Character Holotype
Holotype and 17 paratypes
n Range Mean 6 SD
1. Standard length 136.2 18 88.2–146.7 121.4 6 17.67
2. Body depth 29.4 18 27.2–30.8 29.52 6 0.93
3. Trunk length 84.6 18 82–89.4 86.07 6 1.66
4. Predorsal length 1.1 18 1.1–3.7 1.86 6 0.61
5. Preanal length 19.1 18 17.8–21.3 19.34 6 1.05
6. Dorsal-fin length 98.5 18 94.3–98.6 98.06 6 0.98
7. Anal-fin length 80.9 18 78.8–82.7 81.04 6 1.17
8. Pelvic-fin length 5.9 14 5.9–9.6 7.0 6 0.94
9. Pelvic to anal length 1.8 16 1.2–2.2 1.76 6 0.31
10. Caudal-fin length 8.1 9 7.5–9.5 8.44 6 0.71
11. Head length 16.2 18 13.9–16.2 15.11 6 0.72
12. Head width 25.7 18 21.2–25.7 23.49 6 1.39
13. Postorbital length 11.8 18 9.3–11.8 10.41 6 0.64
14. Upper head lobe width 16.2 18 12.4–16.2 14.47 6 1.04
15. Lower head lobe width 9.6 18 6.6–10.9 8.89 6 1.11
16. Postorbital length 72.7 18 61.9–73.1 68.87 6 3.17
17. Snout length 15.9 18 12.2–23.8 18.02 6 3.35
18. Upper jaw length 18.2 18 18.2–24.4 20.47 6 1.59
19. Eye diameter 6.8 18 5.3–7.9 6.77 6 0.79
20. Chin depth 20.5 18 15.0–26.2 21.75 6 2.56
21. Lower opercular lobe 40.9 18 30.8–40.9 35.49 6 3.03
22. Upper opercular lobe 18.2 18 15–31.8 21.13 6 4.93
346 Copeia 2009, No. 2
Symphurus regan i also has similar dorsal-, anal-, and
caudal-fin rays and similar vertebral counts to those of S.
megasomus, but this species has more abdominal vertebrae
(10, 3+7) and a different ID pattern (1-2-2-2-2). Symphurus
australis, which occurs in deep water off Australia, has more
dorsal-fin rays (111–119), more vertebrae (60–64), and a
different ID pattern (1-2-2-2-2) than that of S. megasomus.
Symphurus fuscus, S. maldivensis, S. ocellatus, S. sayademalhen-
sis, and S. variegatus also share similarities in fin rays and
vertebrae to those of S. megasomus, but these Indian Ocean
species all differ from this species with respect to their 1-2-2-
2-2 ID pattern.
Symphurus monostigmus, from the western Indian Ocean, is
similar to S. megasomus in having 14 caudal-fin rays and a
deep body (22.7–32.8% SL), but differs in having more fin
rays (106–111 dorsal-fin rays, 91–95 anal-fin rays vs. 86
dorsal-fin rays, 73–74 anal-fin rays), a different ID pattern (1-
2-3-2-2 vs. 1-2-2-2-2), in lacking the abdominal pigment
spot that is present in S. monostigmus, and having a larger
body size.
ACKNOWLEDGMENTS
The authors express their appreciation to T. Munroe for help
in improving an early draft of this manuscript and for
providing critical comments and helpful suggestions in
studying tonguefishes. We thank the financial support of
the deep-sea biodiversity project (NSC 90-2311-B-001-121,
NSC 91-2611-B-001-001, and NSC 92-2311-B-001-034) by
National Science Council. H.-M. Yeh, Coastal and Offshore
Resource Research Center, Fisheries Research Institute,
kindly provided information for trawl stations. We also
appreciate the captain and crews of R/V Ocean Researcher I
for their cooperation. Finally, we thank all members of the
Laboratory of Fish Ecology and Evolution, Research Center
for Biodiversity, Academia Sinica for their assistance in
various aspects of this study.
LITERATURE CITED
Chabanaud, P. 1956. Les Symphurus marbre´s du complexe
Indo-Pacifique tropical. Archives du Museum D’Histoire
Naturelle Paris (7
th
Se´ries) 4:79–100.
Gilbert, C. H. 1905. The deep-sea fishes of the Hawaiian
Islands, p. 577–713. In: The Aquatic Resources of the
Hawaiian Islands. D. S. Jordan and B. W. Evermann (eds.).
Bulletin of the United States Fish Commission 23(2).
Heemstra, P. C. 1986. Cynoglossidae, p. 865–868. In:
Smiths’ Sea Fishes. M. M. Smith and P. C. Heemstra
(eds.). Springer-Verlag, Berlin.
Krabbenhoft, T. J., and T. A. Munroe. 2003. Symphurus
bathyspilus: a new cynoglossid flatfish (Pleuronectiformes:
Cynoglossidae) from deepwaters of the Indo-West Pacific.
Copeia 2003:810–817.
Munroe, T. A. 1992. Interdigitation pattern of dorsal-fin
pterygiophores and neural spines, an important diagnos-
tic character for symphurine tonguefishes (Symphurus:
Cynoglossidae: Pleuronectiformes). Bulletin of Marine
Science 50:357–403.
Munroe, T. A. 1998. Systematics and ecology of tongue-
fishes of the genus Symphurus (Cynoglossidae: Pleuronec-
tiformes) from the western Atlantic Ocean. Fishery
Bulletin 96:1–182.
Munroe, T. A. 2006. New Western Indian Ocean tonguefish
(Pleuronectiformes: Cynoglossidae, Symphurus). Copeia
2006:230–234.
Munroe, T. A., and K. Amaoka. 1998. Symphurus hondoensis
Hubbs, 1915, a valid species of Western Pacific tonguefish
(Pleuronectiformes: Cynoglossidae). Ichthyological Re-
search 45:385–391.
Munroe, T. A., and J. Hashimoto. 2008. A new Western
Pacific tonguefish (Pleuronectiformes: Cynoglossidae):
the first pleuronectiform discovered at active hydrother-
mal vents. Zootaxa 1839:43–59.
Ochiai, A. 1984. Cynoglossidae, p. 355–356. In: The Fishes
of the Japanese Archipelago. H. Masuda, K. Amaoka, C.
Araga, T. Uyeno, and T. Yoshino (eds.). Tokai University
Press, Japan.
Shen, S.-C. 1983. Cynoglossid fishes (Pleuronectiformes:
Cynoglossidae) of Taiwan. Bulletin of the Institute of
Zoology, Academia Sinica 22:105–118.
Shen, S.-C. 1984. Coastal Fishes of Taiwan. National Taiwan
University, Taipei.
Shen, S.-C. 1993. Pleuronectiformes, p. 565–585. In: Fishes
of Taiwan. S.-C. Shen, S.-C. Lee, K.-T. Shao, H.-C. Mok,
C.-H. Chen, C.-C. Chen, and C.-S. Tzeng (eds.). National
Taiwan University, Taipei, Taiwan.
Shen, S.-C., and W.-W. Lin. 1984. Some new records of
fishes from Taiwan with descriptions of three new species.
Taiwan Museum Special Publication Series 4:1–25.
Copeia cope-09-02-18.3d 3/3/09 03:56:44 347 Cust # CI-08-080R1
Lee et al.—New species of Symphurus 347
