Symphurus orientalis (Bleeker) redefined based on ...€¦ · ISSN 1175-5326 (print edition) ... Symphurus orientalis (Bleeker) redefined based on morphological and molecular characters

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ISSN 1175-5334 (online edition)Copyright copy 2013 Magnolia Press

Zootaxa 3620 (3) 379ndash403 wwwmapresscomzootaxa Article

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Symphurus orientalis (Bleeker) redefined based on morphological and molecular characters (Pleuronectiformes Cynoglossidae)

MAO-YING LEE1 THOMAS A MUNROE2 amp KWANG-TSAO SHAO3 4

1Department of Aquaculture National Taiwan Ocean University No 2 Peining Rd 20224 Keelung Taiwan E-mail coleopteragmailcom2National Marine Fisheries Service National Systematics Laboratory NOAA Smithsonian Institution PO Box 37012 National Museum of Natural History WCndash57 MRCndash153 Washington DC 20013ndash7012 E-mail munroetsiedu3Laboratory of Fish Ecology and Evolution Biodiversity Research Center Academia Sinica Nankang 11529 Taipei Taiwan E-mail zosktgatesinicaedutw4Corresponding author E-mail zosktgatesinicaedutw

Abstract

Aphoristia (= Symphurus) orientalis Bleeker 1879 collected from an unspecified depth and location in Japanese waters is the first described species of symphurine tonguefish from Indo-Pacific waters The original description with accompanying illustration is based on the unique holotype specimen and provides limited diagnostic characters for this taxon Subsequent to its description the holotype of A orientalis has been lost Limited diagnostic information and loss of the holotype have caused considerable confusion to subsequent systematic studies regarding the identity of this and similar tonguefish species occurring in the Indo-West Pacific region Several often-cited taxonomic accounts purportedly redescribing S orientalis are erroneous because they include more than one species in these redescriptions These erroneous redescriptions not only confused the species concept of S orientalis (Bleeker) but also confounded the systematics of similar Indo-West Pacific tonguefishes Symphurus novemfasciatus Shen and Lin described on two specimens collected in southern Taiwan shares many morphological and pigmentation features similar to those of S orientalis Morphological data from a large series of tonguefishes collected in Taiwanese and Japanese waters as well as molecular data from a smaller number of specimens from these locations including the type locality of S novemfasciatus confirm the presence of only one species S orientalis (Bleeker) among these specimens Symphurus novemfasciatusShen and Lin is therefore regarded as a junior subjective synonym of S orientalis Symphurus orientalis is redefined based on a large series of specimens identified by a consistent set of morphological criteria and a neotype is designated to stabilize nomenclature and systematics of this species Symphurus orientalis differs from congeners by its combination of a predominant 1ndash2ndash2ndash2ndash2 pattern of interdigitation of proximal dorsal-fin pterygiophores and neural spines 12 caudal-fin rays 9 abdominal and 52ndash55 total vertebrae four hypurals 96ndash101 dorsal-fin rays 82ndash89 anal-fin rays 87ndash99 longitudinal scale rows 37ndash42 transverse scales 5ndash11 (usually) distinct complete or incomplete blackish-brown crossbands on the ocular side uniformly white blind side and conspicuous bluish-black peritoneum Documenting morphological variation for S orientalis represents the most important step towards clarification of the identity of this and other symphurine tonguefish species from this region Reliable identification of specimens of S orientalis also provides the foundation for evaluating the status of several other poorly-known nominal species of Indo-West Pacific tonguefishes that have features similar to those of S orientalis Improved identifications will lead to better knowledge on the geographic distribution of S orientalis and these other species as well as to improve estimates of biodiversity and the biogeography of Indo-West Pacific symphurine tonguefishes

Key words Symphurus orientalis Symphurus novemfasciatus flatfish redescription synonym tonguefish species com-plex cryptic species

Introduction and history of the problem

Bleeker (1879) described a new species of tonguefish Aphoristia orientalis based on a single specimen collected from an unspecified location and depth in marine waters off Japan This report represents the first reliable description of a member of Symphurus captured in the Indo-Pacific region [Aphoristia Kaup 1858 is currently

Accepted by J Sparks 10 Jan 2013 published 7 Mar 2013 379

(Eschmeyer 2012) considered a junior subjective synonym of Symphurus Rafinesque 1810 and this generic assignment is followed in this study] Information provided in the original description and derived from the accompanying illustration of S orientalis distinguishes this species as a tonguefish featuring 100 dorsal-fin rays 86 anal-fin rays 12 caudal-fin rays approximately 90 longitudinal scales and 40 transverse scales and with an ocular-side pigmentation pattern consisting of multiple crossbands a uniformly white blind side and black peritoneum

Somewhere in its history the holotype of S orientalis was lost and its whereabouts are unknown (Eschmeyer 2012) Limited diagnostic information contained in the original description compounded by loss of this holotype have rendered it difficult for subsequent researchers interested in the taxonomy and systematics of Indo-West Pacific tonguefishes to adequately distinguish the species described by Bleeker (1879) Of characters discussed in the original description noteworthy is the number of caudal-fin rays (12) reported for the holotype as this is one of the most important characters for differentiating phenetic groups of species within Symphurus (Ginsburg 1951 Chabanaud 1955 Munroe 1992) In an extensive study summarizing information from several thousand specimens of Symphurus Munroe (1992) reported that the number of caudal-fin rays is one of the most conservative and important diagnostic features for identifying species of Symphurus

Several other internal morphological characteristics of symphurine tonguefishes also important in characterizing and diagnosing these species include the pattern of interdigitation of dorsal proximal pterygiophores and neural spines (ID pattern Munroe 1992) and the number of abdominal and total vertebrae (Chabanaud 1955 Munroe 1992) Indo-Pacific species of Symphurus can readily be separated into different phenetic groups based on combinations of the numbers of caudal-fin rays abdominal vertebrae and by their ID patterns (Munroe 1992) Internal features were not reported in the original description of A orientalis and it is not now possible to know what the character states were for this nominal species Thus the species concept of S orientalis as envisioned by Bleeker (1879) is known only from the limited information available in the original description and the accompanying illustration of this nominal species

Reliance on data presented in the original description of S orientalis and the inability to compare the holotype specimen with other nominal species of Symphurus collected subsequently has resulted in considerable taxonomic confusion and misidentifications regarding specimens of Symphurus collected in the Indo-Pacific region Jordan and Starks (1906) for example in their review of the flounders and soles of Japan questioned whether the specimen described by Bleeker actually belonged to Symphurus Okada and Matsubara (1938) and Matsubara (1955) wrote identification keys for fishes occurring in Japanese waters but did not list any catalogue information for specimens they examined In their key to the species of Symphurus they provided ranges for meristic and morphometric features for specimens purportedly of S orientalis However based on specimens we identified as S orientalis (Lee amp Munroe unpubl data) data in Matsubararsquos study overestimates those for S orientalis and appear to be derived from at least two and possibly more sympatric and morphologically similar species of Symphurus featuring 12 caudal-fin rays including some specimens with meristic features much lower than those of S orientalis (Bleeker 1879 Lee amp Munroe unpubl data)

The works of Okada and Matsubara (1938) and Matsubara (1955) had a profound influence on another work that of Ochiai (1959 Japanese edition and English edition 1963) which purportedly redescribed S orientalis Interestingly Ochiairsquos redescription which is based on catalogued specimens reports ranges of several important meristic and morphometric features for this species that are the same as those listed in Matsubararsquos study (1955) It is uncertain if Ochiairsquos work was based on the same specimens as those included in Matsubararsquos study (1955) because catalogue numbers for specimens were not included in the latter study but it seems likely Ochiairsquos study (1959 1963) is the most widely cited work on S orientalis but his redescription includes data from several morphologically similar species with 12 caudal-fin rays A consequence of being so widely cited is that the erroneous data in Ochiairsquos study has been perpetuated in subsequent literature based on this work and this has contributed significantly to the confusion and misunderstanding regarding the range of variation in morphological features in this nominal species

Chen and Weng (1965) based on four specimens potentially represents the first report of S orientalis from Taiwanese waters but according to the number of caudal-fin rays (15) they reported for their specimens it is uncertain whether they actually examined any specimens of this species Specimens included in the study by Chen and Weng (1965) have been lost so it is difficult to know whether these authors counted caudal-fin rays accurately or if their redescription was based on specimens of more than one species at least one of which featured 15 caudal-

LEE ET AL 380 middot Zootaxa 3620 (3) copy 2013 Magnolia Press

fin rays No species of Symphurus described to date has been found to have 15 fin rays as the typical count for caudal-fin rays (Munroe 1992 Lee et al 2009a 2009b) Although some individuals with 15 caudal-fin rays are occasionally encountered in specimens of symphurine species that typically have a count of 14 caudal-fin rays we have never found any specimens with 15 (or even 14) caudal-fin rays among those species examined that characteristically have 12 caudal-fin rays Therefore it seems unlikely that the number of caudal-fin rays was accurately reported in the study of Chen and Weng (1965) and the tonguefishes they examined belonged to species characterized by 14 caudal-fin rays With loss of the specimens their identifications will never be determined

Incorrect information contained in the redescription of S orientalis by Ochiai (1959 1963) has been repeated in much of the subsequent systematic literature dealing with this species (eg Chyung 1961 Amaoka 1982 Shen 1984 Ochiai 1984 Ochiai 1987 Ochiai 1988 Ochiai 1989 Shen 1993 Kim amp Choi 1994 Li amp Wang 1995 Yamada 2000 2002) Several studies dealing with western Pacific Symphurus (Munroe amp Amaoka 1998 Munroe amp Hashimoto 2008) have also noted cases where different nominal species have been incorrectly identified as S orientalis None of these studies however attempted to resolve issues regarding the identity and taxonomic status of S orientalis

Shen and Lin (1984) likely influenced by misinformation on caudal-fin ray counts presented in Chen and Wengrsquos study (1965) for specimens purported to be S orientalis from Taiwanese waters (as evidenced by comparisons section in account of S strictus by Shen 1984141) described the nominal species S novemfasciatus from off Dong-Gang southwestern Taiwan In their description of S novemfasciatus Shen and Lin (1984) did not mention or diagnose their nominal species from the previously described S orientalis despite the fact that it has 12 caudal-fin rays and multiple ocular-side crossbands and also has dorsal- and anal-fin ray counts similar to those features reported for the holotype of S orientalis Subsequent studies recognizing S novemfasciatus as a valid species (Munroe 1992 Shen 1993 Lin 1994 Li amp Wang 1995 Munroe 2000 Liu 2008) have never adequately diagnosed this species from S orientalis or for that matter any of three other previously described Indo-West Pacific species with similar meristic features including S septemstriatus (Alcock 1891) S luzonensis Chabanaud 1955 and S fallax Chabanaud 1957 Consequently the taxonomic status of S novemfasciatus as well as that of these other species is questionable

In recent years symphurine tonguefishes have been collected by commercial fishery trawlers operating off Japan and Taiwan and these fishes are sometimes landed as bycatch species at several different fish ports in these countries Recent collections of fishes taken by research vessels trawling in moderately deep waters off Japan and Taiwan (Lee unpubl data) have also included specimens of Symphurus Based on their relatively frequent occurrence and abundance in both scientific and commercial catches these tonguefishes appear to be more common in deepwater habitats in these waters than previously thought

A large number of the symphurine tonguefishes captured off Japan and Taiwan and preserved in fish collections in these countries have been identified as S orientalis However based on shared similarities in meristic and morphological characteristics it is impossible to distinguish these specimens from S novemfasciatus Most often specimens from Japan have been identified as S orientalis while those taken off southern Taiwan have been identified as either species In addition to this problem among museum lots containing specimens from Taiwanese and Japanese waters that have been identified as S orientalis we have also identified several other diminutive species of tonguefishes featuring 12 caudal-fin rays Taxonomic status of these nominal species is unresolved and currently under study (Lee amp Munroe in prep) However given the overall similarity of these diminutive species to S orientalis it seems likely that specimens of these small-sized species could easily have been misidentified as juvenile S orientalis and included in earlier accounts for that species

As mentioned previously other potential problems regarding the taxonomy of Indo-West Pacific symphurine tonguefishes involve the status of several nominal species similar to S orientalis and S novemfasciatus including S septemstriatus S luzonensis and S fallax These species also have 12 caudal-fin rays and other meristic features similar to those of both S orientalis and S novemfasciatus In the original descriptions of these species none were ever compared and diagnosed from S orientalis Scant information except for that provided in the original descriptions is known about those rarely-caught species which in the case of S luzonensis and S fallax is even more limited because we know these species only from information based on the unique holotype specimens We currently lack reliable information as to how many of these nominal species are valid

Symphurus orientalis (Bleeker 1879) as the oldest available name for an Indo-Pacific species of symphurine tonguefish has priority over any other names proposed subsequently for this species An important first step in

Zootaxa 3620 (3) copy 2013 Magnolia Press middot 381SYMPHURUS ORIENTALIS REDEFINITION

stabilizing the taxonomy and nomenclature for this and other similar nominal species from this region (and beyond) is to more precisely define the concept of S orientalis Proper delineation of this species requires better understanding of the variation present in its morphological features To address this need we provide a redescription of this species based on information from the original description together with new data recovered from examination of a large series of recently-collected specimens that were identified as S orientalis based on their shared similarities with the specimen originally reported in Bleeker (1879) We also designate a neotype for this species to further resolve taxonomic confusion between this and other morphologically similar species

The large number of similarities between S orientalis and S novemfasciatus Shen and Lin 1984 required us to test the hypothesis that specimens from southeastern Taiwan the type locality of S novemfasciatus represent a species distinct from S orientalis from off Japan This comparison was made using both morphological characters and molecular information analyzed from DNA samples of a series of specimens from Taiwan and Japan respectively Recently-developed molecular methods comparing genetic divergence between nominal species can be useful for identifying cryptic species and disentangling problems involving two or more morphologically similar species (Victor 2007 2008 Diaz de Astarloa et al 2008 Pyle et al 2008 Last et al 2010) While these molecular methods can provide evidence for taxonomists to discover cryptic (and often undescribed) species they can also confirm the status of invalid species (Byrkjedal et al 2007 Dooley amp Jimenez 2008) Since molecular approaches do not rely on morphology they provide an independent test to evaluate the status of different populations that share a similar morphology

Because of the confusion surrounding the identity of S orientalis and the status of similar nominal species occurring in the Indo-West Pacific we lack proper understanding of the systematics distribution and biogeography of all of these fishes Tonguefishes purported to be S orientalis have been reported from a variety of locations including that off Vladivostok Russia (Jordan amp Starks 1906 Jordan et al 1913 Okada 1938) in Korean waters (Mori 1928 Mori amp Uchida 1934 Okada 1938 Mori 1952 Kamohara 1958 Ochiai 1959 1963 Chyung 1961 1977 Kim amp Choi 1994 Sakamoto 1997 Yamada 2000 Lin 2001 Youn 2002 Kim et al 2005) off mainland China (Chu 1931 Wu 1932 Fowler 1934 Ochiai 1959 Wang 1993 Li amp Wang 1995 Yamada 2000 Munroe 2000 Lin 2001 Liu 2008) and the Philippine Islands (Fourmanoir 1985) Many of these records are based on species other than S orientalis so it is difficult to know the actual geographic distribution of S orientalis

Resolving the taxonomic status of both S orientalis and S novemfasciatus is an important contribution and first step towards understanding the systematics of these and other nominal species of Symphurus characterized by 12 caudal-fin rays and high counts of meristic features This information in turn provides the foundation for better estimates of the distributions and ecology of these species as well as improving our knowledge concerning the biodiversity and biogeography of Indo-West Pacific tonguefishes

Materials and methods

A total of 94 specimens including preserved specimens in fish collections as well as fresh specimens collected in the field constituted the basis for data collected in this study Included among these specimens were the holotype of S novemfasciatus and 10 other specimens collected from the type locality of S novemfasciatus Tissue samples for molecular analyses involved a muscle biopsy from specimens taken in field collections made in Japan and Taiwan Tissue samples were originally stored in 95 ethanol Voucher specimens for all tissue samples analyzed were subsequently preserved in 10 formalin transferred to 75 ethanol catalogued and deposited in fish collections (see Material examined section) Institutional abbreviations follow those listed in httpwwwasihorgcodonspdf except that of ASIZP (formerly ASIZT) the Biodiversity Research Center Academia Sinica Taipei Taiwan and NMMBP National Museum of Marine Biology and Aquarium Pintung Taiwan Comparative materials for all other Indo-Pacific species of Symphurus included in this study are listed in Munroe (1992) Shen (1993) Munroe and Amaoka (1998) Krabbenhoft and Munroe (2003) Munroe (2006) Munroe and Hashimoto (2008) and Lee et al (2009a 2009b)

Methods for counting meristic characters and for measuring morphometric features and general terminology follow those of Munroe (1998) All 94 specimens examined were radiographed Terminology and formulae for interdigitation patterns of proximal dorsal pterygiophores and neural spines (ID pattern) follow those of Munroe (1992) Morphometric characters were taken to the nearest 01 mm using either dial calipers or a dissecting

microscope fitted with an ocular micrometer Morphometric features are expressed either as proportions in percent standard length (SL) or percent head length (HL)

Description of pigmentation features are based primarily on freshly-landed specimens with supplemental information provided from specimens preserved in formalin and transferred to 75 ethanol Maturity was estimated by macroscopic examination of the extent of posterior elongation 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 are apparent in testis size between mature and immature males therefore estimates of maturity are based entirely on females

To examine genetic divergence in nominal species some researchers (Hebert et al 2003 Ward et al 2005) suggest using sequences of the 5rsquo region of the mitochondrial cytochrome c oxidase subunit I gene (CO I or COX I) for species identification whereas others recommend using 16S rRNA as a good model for assisting taxonomic works (Akimoto et al 2002 Maretto et al 2007 Lakra et al 2009) We chose to examine both genes in an attempt to maximize the information derived from tissue samples Total genomic DNA was extracted from 12 individuals of S orientalis (identifications based on morphological and pigmentation characters consistent with those listed in Bleekerrsquos (1879) original description of this species) including 10 from Taiwan and two from Japan as well as that from 10 individuals tentatively identified as S novemfasciatus (based on their capture at Dong-Gang Taiwan the type locality of this nominal species) For comparative purposes analyses of DNA also included data from three individuals of S strictus Gilbert (identifications based on comparisons with original description) four S hondoensis Hubbs (identifications following redescription of Munroe amp Amaoka (1998)) and six S megasomusLee et al (based on tissue samples of type specimens) DNA was extracted using the Genomic DNA Mini Kit (Geneaid Taipei Taiwan) Approximately 639 base pair (bp) fragments of the COX I gene and 510 bp of the 16S rRNA were amplified using the following primer pair 16Sa-L (5rsquoCGCCTGTTTACCAAAAACATCGCCTrsquo) and 16Sb-H (5rsquoCCGGTCTGAACTCAGATCACGTrsquo) (Palumbi 1996) for the 16S rRNA gene and a newly developed primer pair Symphurus-COIF (5rsquoGGTGCCTGAGCHGGRATAATTGGHACrsquo) and Symphurus-COIR (5rsquoTAAATTTTTGKGTGGCCAAAGAATCArsquo) for the COI gene A polymerase chain reaction (PCR) was carried out using a thermal cycler (BIO-RAD Philadelphia PA USA) in 25-μl reaction volumes containing 100 ng total DNA 1 μM of each primer 04 mM dNTP 1x reaction buffer and 05 U of Taq polymerase (Genomics Taipei Taiwan) with denaturation at 94degC for 4 min followed by 35 cycles of denaturing at 94degC for 30 s annealing at 48degC for 45 s and extension at 72degC for 1 min with a final extension at 72degC for 10 min The PCR products were then sequenced bidirectionally and analyzed on an ABI3730XL model (Applied Biosystems Foster City CA USA)

All sequences were checked against electropherograms and manually edited using the program 4Peaks version 17 (Griekspoor amp Groothuis 2006) In order to confirm the absence of stop codons in the amplified COI we translated the nucleotide sequences with the vertebrate mitochondrial genetic code using EMBOSS-transeq (EMBL-EBI URL httpwwwebiacukToolsstemboss_transeq)

All sequences were aligned with CLUSTAL X version 181 (Thompson et al 1997) Nucleotide genetic distances the Kimura two-parameter distance (K2P) (Kimura 1980) substitution model including transitions and transversions complete deletion of gapsmissing data uniform rates among sites and between and within species comparisons were also calculated using MEGA 40 (Tamura et al 2007) Trees based on sequence data were constructed by the neighbor-joining (NJ) method and evaluated by 10000 bootstrapping replications (Felsenstein 1985) using MEGA 40 (Tamura et al 2007) Trees were constructed only to show divergence in genetic sequences among the samples analyzed We decided to use the neighbor-joining method instead of maximum likelihood and maximum parsimony because our approach in this study focused on species identifications with both COI (DNA barcoding) and 16S rRNA applied for this purpose Sequences were deposited in GenBank (accession numbers JN678732mdashJN678801)

Symphurus orientalis (Bleeker 1879)(Figs1ndash5 Tables 1ndash2)

Aphoristia orientalis Bleeker 1879 31 Pl 2 (fig 1) (Japan description illustration of holotype) Symphurus orientalismdashJordan and Snyder 1901 122 (listed questionable occurrence Japan) Jordan and Starks 1906 243

(synonymy description based on Bleeker (1879) doubted validity of species transfer to Symphurus coasts of Japan north

of Vladivostok based on Schmidt (1904)) Jordan et al 1913 335 (listed in catalogue both coasts of Japan off Vladivostok based on Schmidt (1904)) Hubbs 1915 496 (brief description one specimen Suruga Gulf Japan) Mori 1928 8 (listed Korea) Chu 1931 94 (listed China) Wu 1932 162 (synonymy listed China) Fowler 1934 223 (synonymy redescription Chihli Peking China and Japan figure) Mori and Uchida 1934 33 (listed Korea) Taranetz 1937 148 (in key) Okada 1938 270 (listed Honshu Japan Korea and Vladivostok) Okada and Matsubara 1938 439 (in part brief morphological information for key counts off Japan Pusan and Vladivostok) Chabanaud 1939 27 (listed world catalogue of flatfishes) Mori 1952183 (listed Pusan Korea) Matsubara 1955 1287 (in part) (brief data on meristic and morphometric features Suruga Bay Owase Kochi Japan and Pusan) Kamohara 1958 64 (listed Suruga Bay to Kochi Prefecture Japan and Korea) Ochiai 1959 217 (in part) (redescription based on composite series of specimens meristic and morphometric data following Matsubara (1955) figure 200 m China Sea Yellow Sea and Pacific side of southern Japan) Chyung 1961 657 (in part) (redescription based on Ochiai (1959) Korea) Ochiai 1963 102 (in part) (English edition of Ochiai (1959)) Chen and Weng 1965 102 (in part likely more than one species included in account Tungkong Taiwan brief redescription) Chen 1969 225 226 (in part) (brief description for identification key follows Chen and Weng (1965) figure Dong-Gang Taiwan) Chyung 1977 582 (listed Pusan Korea) Son 1980 (listed east coast of Korea cited from Kim and Choi (1994)) Yasuda et al 1981 19 569 (local name in several languages) Amaoka 1982 302ndash3 408 (redescription based on one specimen color photograph Tosa Bay Japan) Shen 1983 107 (in part) (brief redescription possibly based on composite series of specimens Taiwan) Shen 1984 581 (in part) (redescription based on composite series of specimens figure in key Taiwan) Ochiai 1984 356 (in part) (redescription based on composite series of specimens following that of Ochiai (1959) figure Japan Suruga Bay to Yellow Sea East China Sea) Chen and Yu 1986 830 (in part) (listed brief description for key Taiwan) Shen 1986 264 (Chinese Japanese names) Li 1987 513 (listed figure eastern Yellow Sea to northern South China Sea) Ochiai 1987 931 (in part) (description following Ochiai (1959) color figure Suruga Bay Yellow Sea East China Sea) Ochiai 1988 342 (in part) (Japanese version of Ochiai (1984)) Ochiai 1989 222 (in part) (description following Ochiai (1959) color figure Suruga Bay Yellow Sea East China Sea) Munroe 1992 374 379 (ID pattern meristic information) Lindberg and Federov 1993 207 (mentioned in footnote Japan side sea of Japan) Shen 1993 581 (in part) (redescription based on composite series of specimens black and white photo not this species Taiwan) Wang 1993 115 (listed South China Sea) Kim and Choi 1994810 (no specimens description based on composite series of specimens following Ochiai (1959) Korea (based on Son 1980)) Li and Wang 1995380 (no specimens redescription based on composite series of specimens following Ochiai (1959) illustrations in key China) Sakamoto 1997684 (in part more than one species included in brief account color photo is not S orientalis 200ndash400 m Japan East China Sea Yellow Sea) Munroe and Amaoka 1998 389 (discussed confusion surrounding species concept distinguished from S hondoensis Japan) Eschmeyer 1998 1248 2436 (literature listed as valid species) Evseenko 1998 61 (vertebral count depth of occurrence phylogenetic information of Pleuronectiformes) Schwarzhans 1999366 (description and illustration of otoliths Japan) Yamada 2000 1392 (in part) (brief redescription based on composite series of specimens following Ochiai (1959) in key illustration 200ndash400 m Pacific coast Japan East China Sea Yellow Sea) Munroe 2000 646 (listed South China Sea) Lin 2001 458 (listed Chinarsquos seas including northeastern Yellow Sea to north of South China Sea) Munroe 2001 3895 (listed West Central Pacific) Shinohara et al 2001 337 (listed Tosa Bay Japan) Yoda et al 2002 29 (listed Japanese English names) Yamada 20021392 (in part) (English edition of Yamada (2000)) Youn 2002441 690 (in part) (brief redescription in key Korea) Kim et al (2005) 490 (in part) (brief redescription listed Korea photograph not of this species) Shinohara et al 2005 443 (listed off Ryukyu Islands Japan) Liu 2008 1057 (listed China in eastern part of Yellow Sea and South China Sea off Japan) Munroe and Hashimoto 2008 44 (comments on misidentifications comparisons with S thermophilus) Lee et al 2009b 57 (compared with S multimaculatus) Shen and Wu 2011 763 (brief redescription with illustration Taiwan)

Symphurus arientalis (Bleeker)mdashMinami 1988962 (in part meristic data follows that of Ochiai (1959) description of larval stages Japan)

Symphurus orientulis (Bleeker)mdashLin 1994 747 (listed Chinarsquos seas including northeastern Yellow Sea to north of South China Sea)

Symphurus novemfasciatus Shen and Lin 1984 8 Fig 3 (based on two specimens color photograph Tung-Kong (=Dong-Gang) Taiwan) Shen 1984141 (after Shen and Lin (1984) description color figure Taiwan compared with S septemstriatus (Alcock)) Shen 1986 264 (listed Chinese name) Chen and Yu 1986 831 (listed brief description for key) Munroe 1992 379 (listed in table meristic features following those in original description) Shen 1993 581 (description based on Shen and Lin (1984) color photograph Taiwan) Lin 1994 747 (listed sandy flat southern Taiwan) Li and Wang 1995 383 (no specimens description based on Shen and Lin (1984) black and white photo in key) Eschmeyer 1998 1204 2436 (literature listed as valid species) Munroe 2000 646 (listed South China Sea) Liu 2008 1057 (listed off Dong-Gang southwestern Taiwan) Ho and Shao 201163 (listed type catalogue Taiwan) Shen and Wu 2011 763 (brief description with color photo)

Symphurus cf orientalis (not of Bleeker)mdashSowerby 1930 182 (Symphurus sp listed in Schmidt (1904) likely a species of Cynoglossus) Shen 1984 141 (compared specimen identified as S strictus Gilbert with S orientalis sensu Chen and Weng (1965) Taiwan) Fourmanoir 1985 50 (three specimens Philippine Islands) Hashimoto et al 1988 87 (Kaikata Caldera west of Chichijima Island Ogasawara (Bonin) Islands) Hashimoto et al 1995 585 (hydrothermal vents Minami-Ensei Knoll Mid-Okinawa Trough Western Pacific) Ono et al 1996 223 (hydrothermal vents Kaikata Seamount near Ogasawara (Bonin) Islands South Japan) Fujikura et al 2002 24 (hydrothermal vent Okinawa Trough)

Symphurus orientalis (not of Bleeker) Ohashi and Motomura 2011 115 (brief description from single specimen 70ndash100 m Shibushi Bay Kagoshima)

FIGURE 1 Symphurus orientalis (Bleeker 1879) ASIZP 72344 female 770 mm SL off northeast Taiwan A Ocular-side pigmentation of freshly-caught specimen B Blind-side coloration of same specimen

Neotype BSKU 44238 mature female 910 mm SL Tosa Bay off Kochi Japan bottom trawl 300ndash400 m collected by O Okamura 13 Nov 1987

Counted and measured 91 specimens (547ndash1090 mm SL) Taiwan off northeastern coast ASIZP 72344 mature female 770 mm SL 24ordm4922rsquoN 121ordm5850rsquoE T-W Wang 23 Aug 2007 ASIZP 72345 male 770 mm SL 24ordm5217rsquoN 121ordm5753rsquoE T-W Wang 23 Aug 2007 ASIZP 72346 mature female 817 mm SL 24ordm5217rsquoN 121ordm5753rsquoE T-W Wang 23 Aug 2007 ASIZP 72347 male 771 mm SL 24ordm5217rsquoN 121ordm5753rsquoE T-W Wang 23 Aug 2007 ASIZP 67634 mature female 935 mm SL Nanfang-Ao fish port M-Y Lee 6 Jan 2007 Taiwan off northeastern coast in landings at Da-Shi fish port ASIZP 72340 2 mature females 878ndash910 mm SL M-Y Lee 23 Aug 2007 ASIZP 72372 (JN678742 and JN678777) mature female 729 mm SL M-Y Lee 30 Dec 2009 ASIZP 72373 (JN678743 and JN678778) mature female 860 mm SL M-Y Lee 30 Dec 2009 ASIZP 72374 (JN678744 and JN678779) male 741 mm SL M-Y Lee 30 Dec 2009 ASIZP 72375 (JN678745 and JN678780) mature female 913 mm SL M-Y Lee 30 Dec 2009 ASIZP 72376 (JN678746 and JN678781) male 601 mm SL M-Y Lee 30 Dec 2009 ASIZP 72377 (JN678747 and JN678782) male 640 mm SL M-Y Lee 30 Dec 2009 ASIZP 72378 (JN678748 and JN678783) mature female 888 mm SL M-Y Lee 30 Dec 2009 ASIZP 72379 (JN678749 and JN678784) male 833 mm SL M-Y Lee 30 Dec 2009 ASIZP 72380 (JN678750 and

JN678785) mature female 786 mm SL M-Y Lee 30 Dec 2009 ASIZP 72381 (JN678751 and JN678786) mature female 753 mm SL M-Y Lee 30 Dec 2009 NMMBndashP 1663 male 964 mm SL Y-M Ju 9 Sep 2003 NMMBndashP 6127 mature female 892 mm SL Y-M Ju 8 May 2003 NMMBndashP 6184 male 947 SL Y-M Ju 8 May 2003 NMMBndashP 6186 mature female 856 mm SL Y-M Ju 8 May 2003 NMMBndashP 8631 2 mature females 788ndash890 mm SL T-M Ju 18 Jun 2005 NMMBndashP 9114 10 (2 exam 1 male and 1 immature female) 671ndash782 mm SL Ta-Shi (=Da-Shi) fish port H-W Chen 7 Aug 2008 NMMBndashP 7484 mature female 808 mm SL Y-M Ju 16 Apr 2004 Eastern Taiwan off Su-ao ASIZP 65666 male 677 mm SL 24ordm4775rsquondash24ordm4801rsquoN 122ordm0009rsquondash122ordm0209rsquoE ORE beam trawl 265ndash352 m Fishery Researcher I OCP 273 13 Jun 2005 ASIZP 66767 4 (2 exam females) 669ndash887 mm SL 24ordm5511rsquondash24ordm5747rsquoN 122ordm0473rsquondash122ordm0543rsquoE beam trawl 267ndash430 m Fishery Researcher I CP 291 8 Aug 2005 ASIZP 66821 4 (3 exam male and an immature and mature female) 714ndash831 mm SL 24ordm5707rsquondash24ordm5827rsquoN 122ordm0461rsquondash122ordm0557rsquoE beam trawl 236ndash272 m Fishery Researcher I CP 292 8 Aug 2005 ASIZP 66898 10 (4 exam 2 males 1 immature and 1 mature female) 657ndash794 mm SL 24ordm5570rsquondash24ordm5723rsquoN 122ordm0430rsquondash122ordm0481rsquoE beam trawl 212ndash275 m Ocean Researcher I CP 290 28 Aug 2004 Taiwan off southwestern coast in landings at Dong-Gang fish port ASIZP 67650 male 765 mm SL M-Y Lee 4 Jul 2007 ASIZP 67651 male 838 mm SL M-Y Lee 4 July 2007 ASIZP 67652 male 825 mm SL M-Y Lee 4 Jul 2007 ASIZP 67653 mature female 804 mm SL M-Y Lee 4 Jul 2007 ASIZP 72341 2 males 721ndash811 mm SL M-Y Lee 28 May 2008 ASIZP 72342 6 (4 males and 1 immature and 1 mature female) 691ndash842 mm SL M-Y Lee 28 May 2008 ASIZP 72556 (JN678752 and JN678787) male 791 mm SL M-Y Lee 21 July 2011 ASIZP 72557 (JN678753 and JN678788) immature female 753 mm SL M-Y Lee 21 Jul 2011 ASIZP 72558 (JN678754 and JN678789) immature female 687 mm SL M-Y Lee 21 Jul 2011 ASIZP 72559 (JN678755 and JN678790) immature female 626 mm SL M-Y Lee 21 Jul 2011 ASIZP 72560 (JN678756 and JN678791) male 761 mm SL M-Y Lee 21 Jul 2011 ASIZP 72561 (JN678757 and JN678792) male 628 mm SL M-Y Lee 21 Jul 2011 ASIZP 72562 (JN678758 and JN678793) male 697 mm SL M-Y Lee 21 Jul 2011 ASIZP 72563 (JN678759 and JN678794) male 805 mm SL M-Y Lee 21 Jul 2011 ASIZP 72564 (JN678760 and JN678795) male 619 mm SL M-Y Lee 21 Jul 2011 ASIZP 72565 (JN678761 and JN678796) mature female 723 mm SL M-Y Lee 21 Jul 2011 NMMBndashP 3714 male 721 mm SL Dong-Kang (=Dong-Gang) fish port J-H Wu 2 May 2002 NMMBndashP 5776 2 mature females 772ndash814 mm SL Dong-Kang (=Dong-Gang) fish port Y-M Ju 13 Mar 2003 NMMBndashP 6222 mature female 909 mm SL Tong-Kong (=Dong-Gang) fish port H-C Ho 5 Jul 2007 NMMBndashP 7914 male 859 mm SL Y-M Ju 11 Jun 2004 NMMBndashP 8147 mature female 763 mm SL Y-M Ju 11 Jun 2004 NMMBndashP 6222 3 (2 males and 1 mature female) 619ndash826 mm SL Tong-Kong (=Dong-Gang) fish port C-W Chang 27 Aug 2008 NTUM 04564 Holotype of S novemfasciatus mature female 799 mm SL Tung-kong (=Dong-Gang) S-C Shen 1 Feb 1980 Taiwan off southwestern coast NMMBndashP 7615 2 mature females 654ndash788 mm SL Kao-Hsung Y-M Ju 4 Jul 2004 NMMBndashP 3713 2 mature females 768ndash802 mm SL Fon-Kan fish port 200 m J-H Wu 2 Aug 2001 South China Sea ASIZP 66881 2 (male and mature female) 729ndash778 mm SL Off Siao Liouciou 22ordm2164rsquondash22ordm2229rsquoN 120ordm1155rsquondash120ordm1328rsquoE mini-beam trawl 336ndash395 m Ocean Researcher I PCP 348 9 Mar 2006 Japan Tosa Bay off Kochi in landings at Mimase fish port BSKU 341 mature female 918 mm SL 11 Apr 1951 BSKU 617 male 823 mm SL 5 Feb 1951 BSKU 618 mature female 812 mm SL 5 Feb 1951 BSKU 807 male 925 mm SL 19 Feb 1951 BSKU 808 male 862 mm SL 19 Feb 1951 BSKU 810 male 917 mm SL 19 Feb 1951 BSKU 1585 mature female 963 mm SL 20 Jan 1952 BSKU 3457 mature female 912 mm SL 6 Dec 1953 BSKU 40990 male 704 mm SL off Saga traditional bottom trawl 26 Feb 1985 Tosa Bay off Kochi BSKU 67756 male 585 mm SL RV Kotaka-maru 25 Jul 2003 BSKU 69970 2 (exam 1 male) 547 mm SL 200 m RV Kotaka-maru 7 Oct 2003 BSKU 88734 male 687 mm SL in landings at fish port 3 Mar 2006 Off Murado Cape Kochi BSKU 3528 male 1090 mm SL 1953 Suruga Bay NSMT-P 7471 mature female 973 mm SL 16ndash20 Sep 1968 NSMT-P 49992 male 745 mm SL Off Heda 245ndash440 m 34ordm5974rsquoN 138ordm4517rsquoE 10 Nov 1996 NSMT-P 78389 mature female 863 mm SL 300ndash400 m 1 Apr 1986 USNM 77066 male 564 mm SL 270ndash520 m 16 Oct 1906

FIGURE 2 A Symphurus novemfasciatus Shen and Lin 1984 holotype NTUM 04586 female 799 mm SL off Dong-Gang southwest Taiwan B Blind-side coloration of same specimen C Symphurus orientalis Neotype BSKU 44238 female 910 mm SL collected off Kochi Japan D Blind-side coloration of Neotype

Counted 2 specimens (314ndash338 mm SL) ASIZP 72358 (JN678762 and JN678797) male 338 mm SL off Kochi Tosa Bay Japan 200 m RV Toyohata-maru 17 Jun 2009 ASIZP 72359 (JN678763 and JN678798) male 314 mm SL off Kochi Tosa Bay Japan 200 m RV Toyohata-maru 17 Jun 2009

Diagnosis Symphurus orientalis is distinguished from all congeners by the combination of a predominant 1ndash2ndash2ndash2ndash2 ID pattern 12 caudal-fin rays 9 abdominal vertebrae 52ndash55 total vertebrae four hypurals 96ndash101 dorsal-fin rays 82ndash89 anal-fin rays 87ndash99 longitudinal scale rows 37ndash42 transverse scales 18ndash22 scale rows on the head posterior to the lower orbit and usually with 5ndash11 distinct wide (covering 4ndash8 scales) complete or incomplete dark blackish-brown crossbands on the ocular side an alternating series of rectangular blotches and unpigmented areas (both extending from base to tip of fin) throughout entire lengths of dorsal and anal fins uniformly white blind side and conspicuous bluish-black peritoneum

Description Symphurus orientalis is a medium-sized species reaching sizes to approximately 109 mm SL Meristic characters are summarized in Table 1 Predominant ID pattern 1ndash2ndash2ndash2ndash2 (8391 specimens) Caudal-fin rays 12 (two specimens with 11) Dorsal-fin rays 96ndash101 Anal-fin rays 82ndash89 Pelvic-fin rays 4 Total vertebrae 52ndash55 abdominal vertebrae 9(3 + 6) Hypurals 4 Longitudinal scale rows 87ndash99 Scale rows on head posterior to lower orbit 18ndash22 Transverse scales 37ndash42

TABLE 1 Frequency of meristic characters of Symphurus orientalis Counts for the neotype (BSKU 44238) indicated by an asterisk () those of the holotype of S novemfasciatus (NTUM 04564) indicated by a solid star( )

Proportions of morphometric features are presented in Table 2 Body relatively deep and moderately elongate maximum depth in anterior one-third of body usually at point between anus and fourth anal-fin ray with moderate taper posteriorly from anus to posterior body margin Preanal length smaller than body depth Head moderately short and wide head width slightly shorter than body depth and much greater than head length (HWHL= 105ndash128 = 112) Upper head lobe wider than lower head lobe (UHLLHL= 102ndash151 = 118) slightly

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

Caudal-fin rays Abdominal vertebrae

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

Longitudinal scale count

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

Transverse scale count

37 38 39 40 41 42 N

7 21 21 24 12 1 86

shorter than postorbital length Lower lobe of ocular-side opercle wider than upper opercular lobe posterior margin of lower lobe projecting slightly beyond posterior margin of upper opercular lobe Snout moderately short slightly rounded to obliquely blunt anteriorly its length greater than eye diameter (SNLED= 139ndash211 =162) Dermal

papillae present but not well developed on blind-side snout Ocular-side anterior nostril tubular and short usually not reaching anterior margin of lower eye when depressed posteriorly Ocular-side posterior nostril a small rounded tube located on snout just anterior to interorbital space Blind-side anterior nostril tubular short easily distinguishable from dermal papillae blind-side posterior nostril a shorter and wider posteriorly-directed tube situated posterior to vertical at posterior margin of jaws Jaws long and slightly arched upper jaw length longer than snout length posterior margin of upper jaw usually extending to point between verticals through anterior margin of pupil and midpoint of lower eye Ocular-side lower jaw without fleshy ridge Chin depth slightly shorter than or equal to snout length Eyes moderately large and oval separated by three to four rows of small ctenoid scales in narrow interorbital space Eyes usually equal in position or upper eye slightly in advance of lower eyePupillary operculum absent Dorsal-fin origin located at point between verticals through anterior margin of upper eye and anterior margin of pupil of upper eye predorsal length moderately short Anteriormost dorsal-fin rays slightly shorter than more posterior fin rays Scales absent on both sides of dorsal- and anal-fin rays Pelvic fin moderately long longest pelvic-fin ray when extended posteriorly usually reaching base of first to third anal-fin ray Posteriormost pelvic-fin ray connected to anal fin by delicate membrane (torn in many specimens) Caudal fin relatively long with several rows of ctenoid scales on base of fin Body with numerous strongly ctenoid scales on both sides

TABLE 2 Morphometrics for the neotype (BSKU 44238) and examined specimens of Symphurus orientalis including those for the holotype of S novemfasciatus (NTUM 04564) SL in mm characters 2ndash15 in of SL 16ndash23 in of HL

Character Neotype NTUM 04564 Examined Specimens

n Range Mean plusmn SD

1 Standard length 910 799 92 547ndash1090 7863plusmn1030

2 Body depth 261 268 92 242ndash288 2629plusmn112

3 Trunk length 842 829 90 803ndash851 8293plusmn114

4 Predorsal length 29 36 90 24ndash45 344plusmn042

5 Preanal length 229 225 91 210ndash256 2344plusmn105

6 Dorsal-fin length 970 964 90 948ndash976 9654plusmn047

7 Anal-fin length 773 774 90 745ndash792 7653plusmn104

8 Pelvic-fin length 80 ndash 80 58ndash92 754plusmn089

9 Pelvic to anal length 36 23 84 15ndash48 328plusmn075

10 Caudal-fin length 108 112 72 102ndash127 1130plusmn065

11 Head length 177 197 92 174ndash216 1962plusmn102

12 Head width 200 227 91 190ndash252 2204plusmn129

13 Postorbital length 119 128 91 114ndash147 1318plusmn073

14 Upper head lobe width 108 121 89 102ndash141 1229plusmn082

15 Lower head lobe width 95 111 89 86ndash124 1030plusmn077

18 Snout length 182 174 90 172ndash221 1875plusmn122

19 Upper jaw length 201 205 90 172ndash228 2028plusmn121

20 Eye diameter 117 113 92 97ndash126 1143plusmn064

21 Chin depth 195 163 90 139ndash224 1712plusmn184

22 Lower opercular lobe 292 276 88 218ndash327 2668plusmn226

23 Upper opercular lobe 245 268 88 210ndash314 2565plusmn233

24 HWHL 113 115 91 105ndash128 112plusmn005

25 PupilEye diameter 524 528 92 511ndash771 6181plusmn596

Teeth present and recurved slightly inwards on all jaws but better developed on blind-side jaws Ocular-side premaxilla and dentary with single row of sharply pointed well-developed teeth Blind-side premaxilla with two to four rows of sharp recurved teeth Blind-side lower jaw with three to five rows of well-developed teeth

Coloration of fresh-caught specimens (Fig 1) Body pigmentation generally similar for both sexes at all sizes Ocular-side background coloration generally straw-colored to dark-brown usually with 5ndash11 distinct wide (covering 4ndash8 scales) complete or incomplete dark blackish-brown crossbands crossbands not continued onto dorsal and anal fins some specimens with crossbands faded and indistinct or with uniformly brown pigmentation without crossbands Anteriormost crossband on body region between opercle and vertical through anus successive crossbands present on mid-body region to caudal-fin base External surface of abdominal area usually bluish-black on both sides of body but sometimes with same general coloration as that of ocular-side body (because darker peritoneal pigment obscured by abdominal wall and not visible externally) Background coloration of ocular-side head generally similar to that on body Ocular-side snout light yellow Ocular-side lips and chin region uniformly yellow to brown margins of lips pigmented with small black chromatophores Ocular-side anterior nostril brown Upper aspects of eyes and eye sockets light blue pupils bluish-black Outer surface of ocular-side opercle yellow to brown usually with same background coloration as that of head and body Inner surface of ocular-side opercle and isthmus unpigmented

Blind side generally white to light yellow with bluish-black peritoneum showing through abdominal musculature Outer surface of blind-side opercle uniformly white to light yellowish Inner surface of blind-side opercle unpigmented

Fin rays of dorsal anal and pelvic fins uniformly yellow to brown basal regions of fin rays and membranes covering fin rays light yellow with diffuse scattering of yellow to brown melanophores covering entire fin membranes on both sides of fins Dorsal and anal fins throughout their lengths with alternating series of darkly streaked and lightly pigmented fin rays Basal margins of fin rays and associated fin membranes on blind side light yellow to light brown

Coloration of recently preserved specimens (Fig 2) Similar to that of freshly-caught fishes Specimens stored in preservative for decades usually mostly faded and with only remaining pigmentation consisting of the bluish-black eye sockets and black peritoneum

Size and sexual maturity 94 specimens range in size from 314 to 1090 mm SL Females (n = 49 314ndash1034 mm SL) and males (n = 45 338ndash1090 mm SL) attain similar sizes Nine females (314ndash775 mm SL) are immature and show only slight elongation of the ovaries 40 females ranging in size from 654 to 1034 mm SL are mature with elongate ovaries 29 of these (654ndash1034 mm SL) are mature with elongate ovaries but are non-gravid while another 11 females ranging from 759 to 973 mm SL are gravid

Distribution Symphurus orientalis is known from voucher specimens taken off the continental shelf and upper continental slope including captures in Japanese waters at Suruga Bay Tosa Bay and the Pacific side of southern Japan also in the East China Sea in the Okinawa Trough and from several locations off Taiwan including off I-lan off northeastern and southwestern Taiwan and in the South China Sea off Dong-Gang Taiwan (Fig 3) Based on voucher specimens this species occurs at depths ranging from 200 to 520 m throughout its geographic range with most captures usually occurring between 250 and 400 m Symphurus orientalis is frequently taken on muddy substrata or a mixture of mud-sand substrata

Literature accounts reporting tonguefishes identified as S orientalis list this species from a wider geographic range including the Philippines (Fourmanoir 1985) off mainland China off Korea and off Vladivostok Russia Because of uncertainties involving specimens previously identified as S orientalis reports of S orientalis from some of these locations are questionable

We did not examine the three specimens from the Philippines that Fourmanoir (1985) identified as S orientalis which were taken at similar depths (299ndash320 m) to those occupied by S orientalis Counts he reported for these specimens are at the low ends of ranges of those for S orientalis from Japan and Taiwan respectively Possibly these specimens are S orientalis However they need to be compared with other specimens from the Philippines with 12 caudal-fin rays including those that Chabanaud (1955) identified as S septemstriatus and others representing another nominal species of uncertain status that we (Lee amp Munroe unpubl data) have identified among specimens from the Philippine Islands Both nominal forms are similar to S orientalis but they differ from it in several of their morphometric features

Although several studies (Chu 1931 Sowerby 1930 Wu 1932 Fowler 1934 Li 1987 Lin 1994 Liu 2008) reported the geographic distribution of S orientalis to include waters off mainland China (Bei-Jing and other localities) these records seem doubtful and none are vouchered by specimens Li and Wang (1995) in their work on flatfishes inhabiting Chinese waters commented that reported captures of S orientalis from off mainland China were ldquosuspectrdquo They instead listed the distribution of S orientalis from Taiwan (based on Chen amp Weng 1965) to the eastern part of the Yellow Sea and southern Japan In an extensive study of fishes trawled at depths of 120ndash1100 m in the East China Sea between 26deg and 33degN latitudes Chengyu et al (1986) did not report capturing this species nor have recent collecting efforts off mainland China (X Kong person commun 27 November 2009) collected any Symphurus in the deepwater areas sampled off mainland China Water depths off the coast of China are usually shallower than 100 m and based on what we know concerning depth of capture of specimens of S orientalis from other areas it seems unlikely to find S orientalis in the waters off China

Other studies (Mori 1928 Mori amp Uchida 1934 Son 1980 Kim amp Choi 1994 (based on Son 1980)) record S orientalis from off the east coast of North Korea or from Korean waters (Kim et al 2005) Reported occurrences of S orientalis in waters off the east coasts of North and South Korea are questionable because they are based at least in part on misidentified specimens For example identifications in Kim and Choi (1994) are based on data provided in Ochiairsquos (1959) redescription of S orientalis which includes more than one species (see detailed comments below) Kim et al (2005) in their illustrated book of Korean Fishes record S orientalis from Korean waters and provide a brief description with a color photograph of a specimen purported to be this species However their description of S orientalis follows that of Ochiai (1959) and furthermore the fish in the color photograph that they identify as S orientalis is not that species No voucher specimens were listed in any of the studies (Son 1980 Kim and Choi 1994 Kim et al 2005) reporting S orientalis from Korean waters so it is not now possible to determine if specimens included in those accounts are actually this species Off South Korea and adjacent areas such as the Yellow Sea continental shelf depths are shallower than 150 m These depths are shallower than those typically inhabited by S orientalis (200 m and deeper) based on voucher specimens we examined Most literature (Ochiai 1959 1963 Amaoka 1982 Sakamoto 1997 Yamada 2000 2002 Choi et al 2002) as well as detailed collecting efforts in the Sea of Japan (Yeh 2001 Tian et al 2006) the only known deeper-water area off Korea where S orientalis would most likely be found based on depth of occurrence of voucher specimens we examined have not reported capturing specimens of Symphurus from these waters nor do they include the Sea of Japan as a part of the geographic range of S orientalis Based on the numerous misidentifications of specimens and the lack of documented captures for this species we conclude that reports of S orientalis from Korean waters need confirmation with voucher specimens

Several studies (Jordan amp Starks 1906 Jordan et al 1913 Okada 1938) record S orientalis from off Vladivostok based on Schmidtrsquos (1904) account of Symphurus sp from this area However this locality record for S orientalis is doubtful Jordan and Starks did not examine any specimens of S orientalis including the specimen listed by Schmidt (1904) And reports of S orientalis from this location are based only on the specimen listed in Schmidt (1904) However Schmidtrsquos account of Symphurus sp is problematic because it is based on an 85 mm juvenile specimen in poor condition that is missing most of its scales has damaged fins with several fin rays broken and because the description of this specimen includes so little useful diagnostic information that it can not be unequivocally determined even if it is based on a specimen of Symphurus let alone a specimen that could be positively identified as S orientalis as suggested by its inclusion in the synonymy and distribution sections of Jordan and Starksrsquo (1906) account of S orientalis Even though Schmidt states that his specimen lacks a lateral line which is a diagnostic feature for species belonging to Symphurus it is possible that Schmidt could have examined a juvenile specimen of Cynoglossus that was missing its scales and thus appeared to lack a lateral line(s) For juvenile specimens of some species of Cynoglossus that have lost their scales it is sometimes difficult to determine if they have 0 1 or 2 lateral lines (Jordan amp Starks 1906 Munroe unpubl data) Sowerby (1930) too thought that Schmidtrsquos account of a specimen of Symphurus from off Vladivostok was actually a specimen of Cynoglossus Based on information presented in Schmidtrsquos account of a specimen identified as Symphurus sp we can not conclusively rule out the possibility that Schmidt had a specimen of Symphurus However if Schmidt had actually examined a specimen of Symphurus we can confirm that this specimen belonged to a species other than S orientalis because the reported number of dorsal-fin rays (75) if accurate is well below the range of dorsal-fin rays observed in our specimens of S orientalis (96ndash101 see Table 1) Thus we conclude that reports of S orientalisfrom off Vladivostok are suspect if not erroneous The record of S orientalis from this area is based on the

geographic distribution reported for S orientalis that appeared in Jordan and Starks (1906) This record in turn relied on Schmidtrsquos (1904) report of an unidentified tonguefish specimen that we conclude is not very likely this species if even a specimen of Symphurus Records of symphurine tonguefishes from the continental shelf off Vladivostok need updating based on documented catches and reliable identifications of voucher specimens

Remarks The original description and illustration of the holotype of S orientalis by Bleeker (1879 31) clearly indicate that the holotype (and only specimen) is a symphurine tonguefish characterized by 12 caudal-fin rays and a banded pigmentation pattern At the time of its description S orientalis was differentiated from other described species of Symphurus by its unique combination of number of dorsal- and anal-fin rays scale counts and pigmentation pattern

While the original description of S orientalis contained sufficient information to diagnose this species from other described symphurine tonguefishes known at the time of its discovery information contained in the original description has since proven inadequate to differentiate this from other similar nominal species described or discovered subsequent to Bleekerrsquos study Also because S orientalis was known only from a single specimen the range in values for diagnostically important morphological features were unknown for the species and this uncertainty has undoubtedly contributed to the inadequate redescriptions of this nominal species appearing in works of subsequent ichthyologists Further compounding this difficulty is the subsequent loss of the holotype specimen of S orientalis which has eliminated any possibilities of knowing additional diagnostic characters (ie ID pattern vertebral counts) about Bleekerrsquos nominal species that would facilitate more detailed comparisons with other tonguefishes that have similar features to those reported for the holotype of S orientalis

Matsubararsquos (1955) identification key provided a wide range of meristic counts and morphometric measurements for specimens purported to be S orientalis Ochiairsquos (1959 1963) redescription of S orientalisfollowed the information provided in Matsubara (1955) but Ochiairsquos study is more detailed and also provides catalogue numbers of the examined specimens This is the primary reason that his redescription (Ochiai 1959 1963) of S orientalis has been widely cited as an authoritative work on this species and data from that study have often been repeated in subsequent reports of S orientalis from western Pacific localities However recent discoveries (Lee amp Munroe unpubl data) of several additional nominal species in the western Pacific region that have some similarities to S orientalis reveal that Ochiairsquos redescription of S orientalis likely combined morphological data for this species as well as that for one or more of these other nominal species In continental shelf waters off Japan two nominal species of Symphurus featuring 12 caudal-fin rays (same number as in S orientalis) but with fewer meristic features than those of S orientalis [in fact more similar to those of S microrhynchus (Weber) see Munroe amp Marsh 1997] have recently been discovered (Lee amp Munroe unpubl data) Meristic data for one or both of these species were likely included in the key appearing in Matsubara (1955) and in the redescription of S orientalis by Ochiai (1959 1963) as evidenced by the wide ranges reported for several meristic features observed for these specimens (dorsal-fin rays 86ndash100 anal-fin rays 74ndash86 longitudinal scales 81ndash87) In contrast specimens we identified as S orientalis in our study which represent specimens from localities spanning a wide geographic range including Japan have a much narrower and higher range of values for dorsal- (96ndash101) and anal-fin rays (82ndash89) as well as different counts for longitudinal scales (87ndash99)

Chen and Weng (1965) recorded S orientalis from Taiwanese waters However their report of this species from Taiwan is questionable because their redescription of S orientalis indicates that specimens they identified as S orientalis have 15 caudal-fin rays whereas S orientalis typically has only 12 caudal-fin rays (Bleeker 1879 Munroe 1992 this study) Unfortunately specimens examined by Chen and Weng (1965) are lost precluding possibilities of confirming whether the caudal-fin ray counts reported by Chen and Weng were in error However of the many specimens of Symphurus that we have examined we have not found any specimens of species characterized by having 12 caudal-fin rays that had either 14 or 15 caudal-fin rays We think that specimens examined by Chen and Weng were actually other species of Symphurus such as S bathyspilus Krabbenhoft and Munroe or S multimaculatus Lee et al (2009b) These species which occur in Taiwanese waters (Lee unpubl data) differ from S orientalis in having 14 caudal-fin rays but are similar in their counts of dorsal- and anal-fin-rays

In 1984 Shen and Lin (1984) described another nominal tonguefish species from southern Taiwan S novemfasciatus based on two specimens featuring 12 caudal-fin rays and an ocular-side pigmentation pattern with nine crossbands Perhaps misled by the erroneous caudal-fin ray counts reported earlier in Chen and Weng (1965) for specimens purported to be S orientalis from Taiwanese waters Shen and Lin (1984) did not compare their

specimens with those of Chen and Weng nor did they diagnose or compare their specimens with Bleekerrsquos description of S orientalis from off Japan Neither did they compare their specimens with those included in the redescription of S orientalis by Ochiai (1959) In a later study Shen (1984) mentioned some similarity between S novemfasciatus and the ldquoclosely relatedrdquo S septemstriatus but none of the other subsequent studies dealing with S novemfasciatus (Shen 1993 Lin 1994) compared S novemfasciatus with S orientalis or any of the other previously-named species in this species complex (see further remarks below)

Because of the many similarities between S orientalis and S novemfasciatus the status of this nominal species needed to be evaluated to determine if it is distinct from S orientalis Variations in meristic and morphometric features of 92 specimens of S orientalis collected from different locations ranging from Japan to Dong-Gang Taiwan off northeast Taiwan off Ping-tung southwestern Taiwan including the type locality of S novemfasciatus and from the South China Sea were slight and no clinal trends in morphological variation were evident among these specimens Detailed comparisons of the morphology and pigmentation of the holotype of S novemfasciatus (Fig 2A 2B) with those from this larger series of specimens of S orientalis reveal nearly complete overlap between the two nominal species in many features Several important diagnostic characters such as ID pattern or vertebral counts could not be determined in the holotype of S novemfasciatus because its skeleton has decalcified (precluding observing these features from a radiograph) Meristic features of S novemfasciatus that overlap those of S orientalis include fin-ray (caudal-fin rays 12 in both species dorsal-fin rays 101 vs 96ndash101 in S orientalis anal-fin rays 88 vs 82ndash89 in S orientalis) and scale counts (longitudinal rows 94 vs 87ndash99 in S orientalis transverse scales 39 vs 37ndash42 in S orientalis) Likewise proportions of many morphometric features of the holotype of S novemfasciatus lie within ranges of those features recorded for S orientalis (Table 2) Both nominal species have a relatively deep body (BD of S novemfasciatus = 268 SL vs 242ndash288 in S orientalis) both have their preanal lengths shorter than their body depths (PAL 225 SL vs 21ndash256 in S orientalis) both have a moderately short and wide head with the head width just slightly less than the body depth and much greater than their head length (HWHL = 115 vs 105ndash128 in S orientalis) Other similarities between the two species include the width of the upper head lobe compared with that of the lower head lobe (UHLLHL = 109 in S novemfasciatus vs 102ndash151 in S orientalis) shape and size of their short snouts (SNL 174 HL in S novemfasciatus vs 172ndash221 in S orientalis) and their small eyes (ED 113 HL in S novemfasciatus vs 97ndash126 in S orientalis) Additionally these two nominal species have similar coloration Ocular-side coloration was one feature that Shen and Lin (1984) considered diagnostic for S novemfasciatus among its congeners They considered the nine ocular-side crossbands of S novemfasciatus to be a unique diagnostically important character for this species However Bleeker (1879) had much earlier indicated a banded coloration pattern for S orientalis and we also found that many specimens of S orientalis from across the geographic range of this species including southern Taiwan feature 5ndash11 crossbands on their ocular sides

To gain better understanding of the genetic divergence among tonguefishes that we identified as S orientalis including those from the type locality of S novemfasciatus we compared partial sequences of COI and 16S rRNA between specimens from Japan and northeast Taiwan Genetic divergence among individuals from these widespread locations was shallow for both genes (only 012 K2P distance in the 16S and 022 K2P distance in the COI sequence data) If S novemfasciatus were distinct from S orientalis much greater genetic divergence would be expected between these taxa Ward et al (2005 2009) for example suggested that in fishes an average K2P distance of less than 04 is within the range of variation expected within a species The amount of divergence between S novemfasciatus and S orientalis (16S S novemfasciatus 030 K2P distance S orientalis 013 COI S novemfasciatus 026 K2P distance S orientalis 027) and between the nominal species (16S 021 K2P distance COI 026) is similar to that expected for populations within a species Such shallow genetic divergence observed for populations of S orientalis indicates a lack of structure among these widespread populations (Taiwan and Japan) a finding which further supports the hypothesis that these populations belong to one panmictic species Furthermore the N-J trees (Figs 4ndash5) constructed from both the 16S rRNA and COI sequence data also show this lack of structure between populations representing these two nominal species and also indicate that these individuals are members of the same genetic lineage

Based on nearly complete overlap in morphological features and the low amount of genetic divergence among specimens from Japan and Taiwan all evidence indicates that only one species is represented by this material These data strongly support recognizing S orientalis (Bleeker) with S novemfasciatus as its junior subjective synonym

FIGURE 3 Geographic distribution of Symphurus orientalis based on specimens examined in this study (indicated by triangles) and reliable literature reports (indicated by stars) Questionable localities in the literature where S orientalis has been reported to occur are indicated by a question mark () Symbols may represent more than one locality andor more than a single collection from each locality

FIGURE 4 Neighbor-joining tree (part) from partial 16S rRNA gene sequence of species of Symphurus Numbers above nodes indicate bootstrap values for 10000 replications

FIGURE 5 Neighbor-joining tree (part) from partial COI gene sequence of species of Symphurus Numbers above nodes indicate bootstrap value for 10000 replications

Given the long history of confusion regarding the taxonomic status and identity of S orientalis it is necessary to designate a neotype to stabilize the nomenclature of this nominal species Since the original description of S orientalis (Bleeker) is based on a specimen from Japan the following specimen is designated as the neotype of S orientalis (Bleeker) BSKU 44238 (Figs 2C 2D) 910 mm SL mature (not gravid) female collected by O Okamura Nov 1987 from Tosa Bay off Kochi at a depth between 300 and 400 m Meristic features of the neotype are 1ndash2ndash2ndash2ndash2 ID pattern 12 caudal-fin rays 98 dorsal-fin rays 87 anal-fin rays 9(3+6) abdominal vertebrae 54 total vertebrae 4 hypurals 94 longitudinal scales 39 transverse scales and 20 scale rows on the head from the posterior margin of its lower orbit to the posterior margin of the opercle

Comparisons Among Indo-Pacific Symphurus in addition to S orientalis (including S novemfasciatus) five other named species also feature 12 caudal-fin rays (Alcock 1891 Alcock 1896 Weber 1913 Chabanaud 1955 Chabanaud 1957) Of these only S septemstriatus S luzonensis and S fallax have similar fin-ray andor vertebral counts to those of S orientalis

Symphurus septemstriatus is known from relatively few specimens collected in deep waters (260ndash730 m) of the Indian Ocean including the Andaman Sea the Laccadive Sea off Colombo Sri Lanka and in the Gulf of Mannar (Alcock 1891 1896 and 1899 respectively) Opportunities to directly study the types and other specimens of this species curated at the Zoological Survey of India were not available to us Nor is there any published information on morphological features of this species for any other specimens from Indian Ocean localities beyond counts and measurements of the holotype (Norman 1928 Munro 1955) Records of this species and accompanying morphological data from specimens collected at localities beyond these Indian Ocean locations (eg the Philippines in Chabanaud (1955)) may be that for S septemstriatus but these specimens require further study to determine their identity (Lee amp Munroe unpubl data) We do however have a photograph of the holotype of Aphoristia septemstriata which provides some useful comparative information on this species Based on our data S orientalis differs from S septemstriatus in having more anal-fin rays (82ndash89 vs 80 in S septemstriatus) and its upper head lobe is larger than the lower head lobe (UHLLHL = 102ndash151 in S orientalis vs UHLLHL lt 10 in S septemstriatus) The snout of S orientalis is rounded or obliquely blunt anteriorly versus pointed and narrower in S septemstriatus and the migrated eye has a more medial position compared with that of S septemstriatus which has the migrated eye located closer to the dorsal margin of the head Shen (1984) had noted a difference in number of ocular-side crossbands between his S novemfasciatus (= S orientalis) and S septemstriatus however when a larger series of S orientalis are examined this apparent difference does not exist as the two species overlap completely in this feature

Symphurus luzonensis is another nominal species of western Pacific deepwater tonguefish described from a single specimen that was collected off Luzon Island Philippines (Chabanaud 1955) Chabanaud (1955) reported that this specimen had 10 abdominal vertebrae and 52 total vertebrae and thus only diagnosed his species from S regani Weber a species that has 10 abdominal vertebrae and a similar number of total vertebrae A radiograph of the holotype of S luzonensis however reveals that this species actually has 9 abdominal and 53 total vertebrae and an ID pattern and fin-ray counts that are more similar to those of S orientalis (Munroe 1992) Symphurus orientalis differs from S luzonensis in having more scales in a transverse row (37ndash42 vs 34 in S luzonensis) a deeper body (BD 242ndash288 SL vs 221 in S luzonensis) a wider upper opercular lobe (OPUL 210ndash314 HL vs 189 in S luzonensis) larger HWHL ratio (HWHL = 105ndash128 vs 099 in S luzonensis) and its dorsal-fin origin is located more anteriorly than is that of S luzonensis

Symphurus fallax Chabanaud is another poorly-known nominal species described from a small (45 mm SL) damaged holotype specimen which was collected at 397 m off Kei Island Indonesia (Weber 1913) No photograph or illustration accompanied the original description of this nominal species (Chabanaud 1957) nor was any provided in Weberrsquos (1913) or Weber and de Beaufortrsquos (1929) accounts of the specimen which they referred to as an unidentified species of Aphoristia or Symphurus respectively Further compounding problems with the identity of this nominal species is that Chabanaud did not return the holotype to the Zoological Museum of Amsterdamrsquos fish collection (Eschmeyer 2012) Thus the only information on this species is that contained in the original description Of interest is that in the description Chabanaud did not differentiate his nominal species from S orientalis despite the fact that based on our assessments his species was morphologically similar to this congener especially with respect to the numbers of caudal-fin rays We found only minor differences in meristic features between S orientalis and those reported for S fallax (dorsal-fin rays 96ndash101 in S orientalis vs 95 in S fallax) and S orientalis has a slightly deeper body (BD 242ndash288 SL vs 220 in S fallax) Otherwise little

else differentiates these nominal species a finding which questions the validity of S fallax Before arriving at a taxonomic decision regarding the status of this nominal species more specimens from the type locality are needed to determine if S fallax is valid and distinct from S orientalis and S septemstriatus

Discussion

Most problems regarding the taxonomic status of many nominal deep-water species of Symphurus occurring in Indo-Pacific waters are due to the limited amount and overall poor condition of many of the study specimens incomplete and insufficient descriptions and by an overall similarity in phenetic features among several of these nominal species (Munroe 1992 Munroe amp Amaoka 1998 Munroe amp Hashimoto 2008) Symphurus orientalis is now known from a comparatively large number (over 100) of preserved specimens identified by a consistent set of diagnostic features Results of this study provide robust morphological data a detailed description and updated information on the geographic and bathymetric distributions of this species

This study also resolves the taxonomic status of S novemfasciatus Shen and Lin as it was shown that this nominal species is invalid based on the nearly complete overlap in morphological and pigmentation features between it and S orientalis In recent years molecular tools (Tautz et al 2003) including DNA barcoding (Hebert et al 2003) and knowledge about DNA sequences have become more common in helping to resolve taxonomic problems involving closely-related species With respect to most taxonomic groups including some tonguefishes (Tunnicliffe et al 2010) studies of DNA sequences have revealed that estimates of biodiversity at the species level that are based on morphological characters actually underestimate the real diversity represented among the specimens (see also Hebert et al 2004 Ward et al 2008 Teletchea 2009 Tornabene et al 2010) Molecular tools have been especially valuable in discovering cryptic species present among specimens previously considered to represent only a single nominal species Cryptic species show only minor differences in morphology and these minor differences are largely ignored as insignificant features useful for identification Among tonguefishes one example of cryptic species was found among allopatric populations occurring on seamounts in the Pacific Ocean when Tunnicliffe et al (2010) observed large genetic divergence in partial 16S rRNA (90) and COI (142) gene sequences between populations that had previously (Munroe amp Hashimoto 2008) been considered as one species S thermophilus

In light of these findings we tested the hypothesis that two nominal species might be represented among specimens we studied S orientalis which inhabits marine waters from Japan to the Pacific side of Taiwan and a second cryptic species S novemfasciatus with a more limited and allopatric distribution in the region of Dong-Gang South China Sea This hypothesis seemed plausible because these nominal species share so many similarities in their morphological features and pigmentation but are geographically separated The slight genetic divergences we found between populations hypothesized to represent S novemfasciatus and S orientalis both in the partial 16S (021 K2P distance) and COI (026 K2P distance) gene sequences indicated that S novemfasciatus should not be considered a cryptic species Rather based on these results close genetic similarity among these geographically separated populations (Taiwan and Japan) supports conclusions based on morphological data that S novemfasciatus is a junior subjective synonym of S orientalis

Despite improvements in our understanding of morphological and genetic variation in S orientalis a large need still remains for more reliable information on several important biological aspects of this species including its geographic and bathymetric distributions diet reproduction life history demographics microhabitat preferences population density and spatial distribution Better definition of the species concept of S orientalis and an improved understanding of the morphological variation within populations of this species should lead to more reliable identification of specimens of this and related species in the future In turn this should lead to improvements in data documenting its bathymetric and geographic distributions and should provide better insight into the ecology of this species

From a systematics perspective on the genus Symphurus now that the status of S orientalis and S novemfasciatus has been critically evaluated and resolved the task of validating the status of similar nominal species and other populations of Indo-West Pacific Symphurus can begin Questions still abound considering the status and identity of S luzonensis and S fallax which are known only from holotype specimens It remains to be determined how many other Indo-West Pacific species together with S orientalis and S septemstriatus compose

the species complex whose members are characterized by a 1ndash2ndash2ndash2ndash2 ID pattern 12 caudal-fin rays and high meristic features Further expeditions both to locations where specimens of rare and poorly-known species were originally collected and also to those areas that have never been sampled are needed to provide additional specimens to resolve the systematic problems surrounding these tonguefishes

Acknowledgments

This work represents a portion of a collecting activities grant investigating the biodiversity and systematics of deep-sea fishes in Taiwanese waters supported by the National Science Council (NSC 96ndash2628ndashBndash001ndash006ndashMY3 and NSC 99ndash2621ndashBndash001ndash008ndashMY3) and awarded to K-T Shao Biodiversity Research Center Academia Sinica We thank H-M Yeh Coastal and Offshore Resource Research Center Fisheries Research Institute who kindly provided trawl station data from research vessels H-C Ho P-F Lee Y-C Liao and L-P Lin assisted with collecting specimens and kindly provided other specimens used in this study K-C Hsu advised M-Y Lee regarding molecular aspects of this research study A Collins NSL kindly provided information and helpful editorial suggestions regarding molecular approaches used in this study L-P Lin and C-W Chang assisted with loans and shipments of specimens M Nizinski provided assistance and support during M-Y Leersquos visit to the National Museum of Natural History Smithsonian Institution H Endo N Nakayama and R Asaoka provided assistance and support during M-Y Leersquos visit to the Department of Biology Faculty of Science Kochi University They also provided two important fresh specimens of S orientalis from Japan to help with molecular comparisons K Matsuura and G Shinohara provided loan of Suruga Bay specimens during M-Y Leersquos visit to the Department of Zoology National Science Museum K Murphy (USNM) assisted with specimen and catalogue information K Nakaya Hokkaido University provided the photograph of the holotype of S septemstriatus K Vinnikov University of Hawaii at Manoa graciously provided translations of Russian literature L Willis NMFS-NSL assisted with literature H-M Chen provided comments on an early draft of the manuscript M-Y Lee extends his appreciation to all members of the Laboratory of Fish Ecology and Evolution for their support especially H Lee for helping to draw the distribution map and assistance during this study

Literature cited

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Alcock AW (1891) Class Pisces In II ndashNatural history notes from H M Indian marine survey steamer lsquoInvestigatorrsquo Commander R F Hoskyn R N commanding Series II No 1 On the results of deep-sea dredging during the season 1890ndash91 The Annals and Magazine of Natural History (Series 6) 8 (4344) 119ndash138

Alcock AW (1896) Natural history notes from H M Indian marine survey steamer lsquoInvestigatorrsquo Commander C F Oldham R N commanding Series II No 23 A supplementary list of the marine fishes of India with description of 2 new genera and eight new species Journal of the Asiatic Society of Bengal Volume 65 (part II) Number 3 301ndash338

Alcock AW (1899) A descriptive catalogue of the Indian deep-sea fishes in the Indian Museum collected by the RIMSInvestigator Government Printing Calcutta 220 pp

Amaoka K (1982) Symphurus orientalis In Okamura O Amaoka K amp Mitani F (Eds) Fishes of the Kyushu-Palau Ridge and Tosa Bay The intensive research of unexploited fishery resources on continental slopes Japan Fisheries Resource Conservation Association Tosho Printing Co Tokyo pp 303 408 (Japanese and English)

Bleeker P (1879) Eacutenumeraacutetion des espegraveces de poissons actuellement connues du Japon et description de trois espegraveces ineacutedites Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen Afdeling Natuurkunde 18 1ndash33

Byrkjedal I Rees DJ amp Willassen E (2007) Lumping lumpsuckers molecular and morphological insights into the taxonomic status of Eumicrotremus spinosus (Fabricius 1776) and Eumicrotremus eggvinii Koefoed 1956 (Teleostei Cyclopteridae) Journal of Fish Biology 71(suppl A) 111ndash131 httpdxdoiorg101111j1095-8649200701550x

Chabanaud P (1939) Catalogue systeacutematique et chorologique des Teacuteleacuteosteacuteens dyssymeacutetriques du globe Bulletin de LrsquoInstitut Oceacuteanographique (Foundation Albert Ier Prince de Monaco) Number 763 31 pp

Chabanaud P (1955) Flatfishes of the genus Symphurus from the USS ldquoAlbatrossrdquo Expedition to the Philippines 1907ndash1910 Journal of the Washington Academy of Sciences 45(1) 30ndash32

Chabanaud P (1957) Description dun Symphurus ineacutedit fruit de la croisiegravere 1899ndash1900 du Siboga Beaufortia 5(62) 183ndash185

Chen JTF (1969) A synopsis of the vertebrates of Taiwan Revised edition Volume 1 The Commercial Press Taipei 548 pp (in Chinese)

Chen JTF amp Weng HTC (1965) A review of the flatfishes of Taiwan Biological Bulletin Tunghai University Ichthyology Series (5) 25 amp 27 1ndash103

Chen JTF amp Yu MJ (1986) A synopsis of the vertebrates of Taiwan Revised and enlarged edition Volume 2 The Commercial Press Taipei 443ndash1092 pp (in Chinese)

Chengyu X Siming D Hongxi Z amp Keqin H (1986) Studies on the fish fauna of the outer region of the continental shelf and the continental slope in the East China Sea Transactions of the Chinese Ichthyological Society (5) 47ndash63 (Chinese with English abstract)

Choi Y Kim J-H amp Park JY (2002) Marine fishes of Korea Kyo-Hak Publishing Seoul 647 pp (In Korean)Chu YT (1931) Index Piscium Sinensium Biological Bulletin of St Johnrsquos University Shanghai Number 1 291 ppChyung MK (1961) Illustrated Encyclopedia The fauna of Korea (2) Fishes Iljiisa Seoul 861 pp (In Korean)Chyung MK (1977) The fishes of Korea Iljiisa Seoul 727 pp (In Korean)Diaz de Astarloa JM Mabragana E Hanner R amp Figueroa DE (2008) Morphological and molecular evidence for a new

species of longnose skate (Rajiformes Rajidae Dipturus) from Argentinean waters Zootaxa 1921 35ndash46Dooley JK amp Jimenez L (2008) Hoplolatilus luteus Allen amp Kuiter 1989 a junior synonym of H fourmanoiri Smith1964

(Perciformes Malacanthidae) based on morphological and molecular data Aqua International Journal of Ichthyology 14(2) 77ndash84

Eschmeyer WN (1998) Catalog of Fishes Center for Biodiversity Research and Information Special Publication 1 California Academy of Sciences San Francisco 3 Volumes 2905 pp

Eschmeyer WN amp Fricke R (Eds) (2012) Catalog of Fishes Electronic versionndash15 March 2012 California Academy of Sciences Avalaible from httpresearchcalacademyorgichthyologycatalogfishcatmainasp (March 15 2012)

Evseenko SA (1998) The family Achiropsettidae and its position in taxonomical and ecological classifications of the order Pleuronectiformes Russian Academy of Science 62 pp

Felsenstein J (1985) Confidence limits on phylogenies an approach using bootstrap Evolution 39 783ndash791 httpdxdoiorg1023072408678

Fourmanoir P (1985) Poissons Liste et description de cinq espegraveces nouvelles (MUSORSTOM II) Fishes List and description of five new species (MURSORSTOM II) Reacutesultats des campagnes Musorstom I et II Philippines (1976 1980) Tome 2 Meacutemoires du Museacuteum National drsquoHistoire Naturelle Nouvelle Seacuterie Seacuterie A Zoologie Tome 133 31ndash54

Fowler HW (1934) A synopsis of the fishes of China Part V continued Hong Kong Naturalist 5(3) 210ndash225Fujikura K Hashimoto J amp Okutani T (2002) Estimated population densities of megafauna in two chemosynthesis based

communities a cold seep in Sagami Bay and a hydrothermal vent in the Okinawa Trough Benthos Research 57(1) 21ndash30

Ginsburg I (1951) Western Atlantic tonguefishes with descriptions of six new species Zoologica (New York) 36 185ndash201Griekspoor A amp Groothuis T (2006) 4Peaks version 17 Computer program distributed by the authors Avalaible from

httpwwwmekentosjcomscience4peaksHashimoto J Ohta S Fujikura K amp Miura T (1995) Microdistribution pattern and biogeography of the hydrothermal vent

communities of the Minami-Ensei Knoll in the Mid-Okinawa Trough Western Pacific Deep-Sea Research Part I 42(4) 577ndash598 httpdxdoiorg1010160967-0637(94)00037-S

Hashimoto J Hotta H amp Deep Sea Survey Group (1988) The biological communities found at the vicinity of the central cone of the Kaikata Caldera west of Chichijima Island Ogasawara (Bonin) Islands Abstract Papers 5th symposium of Shinkai 2000 study JAMSTEC Journal of Deep Sea Research 5 87ndash88 (in Japanese)

Hebert PDN Cywinska A Ball SL amp de Waard JR (2003) Biological identifications through DNA barcodes Proceedings of the Royal Society B 270(1512) 313ndash322 httpdxdoiorg101098rspb20022218

Hebert PDN Penton EH Burns JM Janzen DH amp Hallwachs W (2004) Ten species in one DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator Proceeding of the National Academy of Sciences of Philadelphia 101(41) 14812ndash14817

Ho H-C amp Shao K-T (2011) Annotated checklist and type catalog of fish genera and species described from Taiwan Zootaxa Number 2957 1ndash74

Hubbs C (1915) Flounders and soles from Japan collected by United States Bureau of Fisheries Steamer ldquoAlbatrossrdquo in 1906 Proceedings of the United States National Museum 48(2082) 449ndash496 httpdxdoiorg105479si0096380148-2082449

Jordan DS amp Snyder JO (1901) A preliminary check-list of the fishes of Japan Annotationes Zoologicae Japonensis 3 1ndash159

Jordan DS amp Starks EC (1906) A review of the flounders and soles of Japan Proceedings of the United States National Museum 31(1484) 161ndash246 httpdxdoiorg105479si0096380131-1484161

Jordan DS Tanaka S amp Snyder JO (1913) A catalogue of the fishes of Japan Journal of the College of Science Imperial University Tokyo 33(1) 1ndash497

Kamohara T (1958) A catalogue of fishes of Kochi Prefecture (Province Tosa) Japan Report of the Usa Marine Biological Station 5(1) 1ndash76 (English with Japanese names)

Kim I-S amp Choi Y (1994) A taxonomic revision of the family Cynoglossidae (Pisces Pleuronectiformes) from Korea

Bulletin of the Korean Fisheries Society 27(6) 803ndash813Kim I-S Choi Y Lee C-Y Lee Y-J Kim B-J amp Kim J-H (2005) Illustrated booked of Korean Fishes Kyohak

Publishing Company Limited 615 pp (In Korean)Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of

nucleotide sequences Journal of Molecular Evolution 16 111ndash120 httpdxdoiorg101007BF01731581Krabbenhoft TJ amp Munroe TA (2003) Symphurus bathyspilus a new cynoglossid flatfish (Pleuronectiformes

Cynoglossidae) from deepwaters of the Indo-West Pacific Copeia 2003(4) 810ndash817 httpdxdoiorg101643IA02-1891

Lakra WS Goswami M amp Gopalakrishnan A (2009) Molecular identification and phylogenetic relationships of seven Indian sciaenids (Pisces Perciformes Sciaenidae) based on 16S rRNA and cytochrome c oxidase subunit I mitochondrial genes Molecular Biology Reports 36(5) 831ndash839 httpdxdoiorg101007s11033-008-9252-1

Last PR White WT amp Puckridge M (2010) Neotrygon ningalooensis n sp (Myliobatoidei Dasyatidae) a new maskray from Australia Aqua International Journal of Ichthyology 16(2) 37ndash50

Lee M-Y Shao K-T amp Chen H-M (2009a) A new species of deep-water tonguefish Genus Symphurus (Pleuronectiformes Cynoglossidae) from Taiwan Copeia 2009(2) 342ndash347 httpdxdoiorg101643CI-08-080

Lee M-Y Munroe TA amp Chen H-M (2009b) A new species of tonguefish (Pleuronectiformes Cynoglossidae) from Taiwanese waters Zootaxa 2203 49ndash58

Li S-Z (1987) Pleuronectiformes In Cheng Q amp Zheng B (Eds) Systematic synopsis of Chinese fishes Volume I Science Press Beijing pp 489ndash513 (In Chinese)

Li S-Z amp Wang H-M (1995) Fauna Sinica Osteichthyes Pleuronectiformes Science Press Beijing 433 pp (Chinese with English summary)

Lin S-D (with edits by Qiu S-Y Yang J-J Lin F-S amp Shao K-T) (1994) Vertebrata In Huang Z-G (Ed) Marine species and their distributions in Chinarsquos Seas China Ocean Press Beijing pp 658ndash764 (In Chinese)

Lin S-D (with edits by Qiu S-Y Yang J-J Lin F-S amp Shao K-T) (2001) Vertebrata In Huang Z-G (Ed) Marine species and their distribution in Chinarsquos Seas Krieger Publishing Company Malabar Florida pp 404ndash463 (English edition)

Lindberg GU amp Federov VV (1993) Fishes of the sea of Japan and adjacent parts of the Sea of Okhotsk and the Yellow Sea Part 6 Teleostomi Osteichthyes Actinopterygii XXXI Pleuronectiformes (CXCV Fam Psettodidae mdashCCI Fam Cynoglossidae) Handbook on the Identification of Animals Zoological Institute of the Russian Academy (166) 1ndash271 (In Russian)

Liu J (2008) Cynoglossidae In Liu R (Ed) Checklist of Marine Biota of China Seas Science Press Beijing pp 1056ndash1057 (Chinese and English)

Maretto F Reffo E Dalvit C Barcaccia G amp Mantovani R (2007) Finding 16S rRNA gene-based SNPs for the genetic traceability of commercial species belonging to Gadiformes Italian Journal of Animal Science 6(1) 161ndash163

Matsubara K (1955) Fish Morphology and Hierarchy Ishizaki Shoten Tokyo Part II pp 790ndash1605 (In Japanese)Minami T (1988) Family Cynoglossidae In Okiyama M (Ed) An atlas of the early stage fishes in Japan Tokai University

Press pp 959ndash964 (In Japanese)Mori T (1928) A catalogue of the fishes of Korea Journal of the Pan-Pacific Research Institution 3(3) 2ndash8Mori T (1952) Checklist of the fishes of Korea Memoirs of the Hyogo University of Agriculture 1(3) 1ndash228Mori T amp Uchida K (1934) A revised catalogue of the fishes of Korea Journal of the Chosen Natural History Society

Number 19 12ndash33 (In Japanese)Munro ISR (1955) The marine and fresh water fishes of Ceylon Department of External Affairs Canberra Australia 351

ppMunroe TA (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) 357ndash403

Munroe TA (1998) Systematics and ecology of tonguefishes of the genus Symphurus (Cynoglossidae Pleuronectiformes) from the western Atlantic Ocean Fishery Bulletin 96(1) 1ndash182

Munroe TA (2000) Family Cynoglossidae In Randall JE amp Lim KKP (Eds) A checklist of the fishes of the South China Sea Raffles Bulletin of Zoology Supplement Number 8 The Biodiversity of the South China Sea pp 646ndash647

Munroe TA (2001) Family Cynoglossidae In Carpenter KE amp Niem VH (Eds) Species identification guide for fishery purposes The living marine resources of the western central pacific Bony fishes part 4 Food and Agriculture Organization of the United Nations Rome pp 3890ndash3901

Munroe TA (2006) New western Indian Ocean tonguefish (Pleuronectiformes Cynoglossidae Symphurus) Copeia 2006(2) 230ndash234 httpdxdoiorg1016430045-8511(2006)6[230NWIOTP]20CO2

Munroe TA amp Amaoka K (1998) Symphurus hondoensis Hubbs 1915 (Cynoglossidae Pleuronectiformes) a valid species of western Pacific tonguefish Ichthyological Research 45(4) 385ndash391 httpdxdoiorg101007BF02725191

Munroe TA amp Hashimoto J (2008) A new Western Pacific Tonguefish (Pleuronectiformes Cynoglossidae) The first pleuronectiform discovered at active hydrothermal vents Zootaxa 1839 43ndash59

Munroe TA amp Marsh BN (1997) Taxonomic status of three nominal species of Indo-Pacific symphurine tonguefishes (Symphurus Cynoglossidae Pleuronectiformes) Ichthyological Research 44(2) 189ndash200 httpdxdoiorg101007

BF02678697Norman JR (1928) The flatfishes (Heterosomata) of India with a list of the specimens in the Indian Museum Part II Records

of the Indian Museum 30 173ndash215Ochiai A (1959) Morphology taxonomy and ecology of the soles of Japan Laboratory of Fisheries Faculty of Science Kyoto

University Kyoto 236 pp (In Japanese)Ochiai A (1963) Fauna Japonica Soleina (Pisces) Biogeographical Society of Japan Tokyo 114 ppOchiai A (1984) Family Cynoglossidae In Masuda H Amaoka K Araga C Uyeno T amp Yoshino T (Eds) The fishes of

the Japanese Archipelago Tokai University Press Tokyo pp 355ndash356 Ochiai A (1987) Family Cynoglossidae In Abe T (Ed) Illustrated fishes of the world in colour Hokuryukan Company

Limited Tokyo pp 927ndash931 (In Japanese)Ochiai A (1988) Family Cynoglossidae In Masuda H Amaoka K Araga C Uyeno T amp Yoshino T (Eds) The Fishes of

the Japanese Archipelago (2nd Edition) Tokai University Press Tokyo pp 341ndash342 (In Japanese)Ochiai A (1989) Cynoglossidae In Abe T amp Ochiai A (Eds) Keys to the Japanese fishes fully illustrated in color

Hokuryukan Company Limited Tokyo pp 219ndash222 (In Japanese)Ohashi Y amp Motomura H (2011) Pleuronectiform fishes of northern Kagoshima prefecture Japan Nature of Kagoshima 37

71ndash118Okada Y (1938) A catalogue of vertebrates of Japan Maruzen Company Limited Tokyo 412 pp (English and Japanese)Okada Y amp Matsubara K (1938) Keys to the fishes and fish-like animals of Japan including Kuril Islands Southern

Sakhalin Bonin Islands Ryukyu Islands Korea and Formosa Sanseido Company Limited Tokyo 584 pp (In Japanese)Ono T Fujikura K Hashimoto J Fujiwara Y amp Segawa S (1996) The hydrothermal vent community at the Kaikata

Seamount near Ogasawara (Bonin) Islands South Japan JAMSTEC Journal of Deep Sea Research 12 221ndash230 (In Japanese with English abstract and legends)

Palumbi SR (1996) Nucleic acids II the polymerase chain reaction In Hillis DM Moritz C amp Mable BK (Eds) Molecular systematics 2nd Edition Sinauer Associates Inc Sutherland MA pp 205ndash247

Pyle R Earle JL amp Greene BD (2008) Five new species of the damselfish genus Chromis (Perciformes Labroidei Pomacentridae) from deep coral reefs in the tropical western Pacific Zootaxa 1671 3ndash31

Sakamoto K (1997) Pleuronectidae Soleidae and Cynoglossidae In Okamura O amp Amaoka K (Eds) Sea fishes of Japan Yama-kei Publishing Company Limited Tokyo pp 672ndash684 (In Japanese)

Schmidt PJ (1904) Fishes of the eastern seas of the Russian empire Scientific results of the Korea-Sakhalin Expedition of the Emperor Russian Geographical Society 1900ndash1901 Saint Petersburg 466 pp (In Russian)

Schwarzhans W (1999) A comparative morphological treatise of recent and fossil otoliths of the order Pleuronectiformes Volume 2 Piscium catalogus Pt Otolithi piscium (Pfeil FH Ed) Muumlnchen Germany 392 pp

Shen S-C (1983) Cynoglossid fishes (Pleuronectiformes Cynoglossidae) of Taiwan Bulletin of the Institute of Zoology Academia Sinica 22(1) 105ndash118

Shen S-C (1984) Coastal fishes of Taiwan Shih-Chieh Shen Department of Zoology National Taiwan University Taipei 190 pp

Shen S-C (1986) Chinese-English-Japanese scientific glossary of fishes Shih-Chieh Shen Taiwan Museum Taipei 427 ppShen S-C (1993) Pleuronectiformes In Shen S-C Lee S-C Shao K-T Mok H-C Chen C-H Chen C-C amp Tzeng

C-S (Eds) Fishes of Taiwan National Taiwan University Taipei Taiwan pp 565ndash585 (In Chinese) Shen S-C amp Lin W-W (1984) Some new records of fishes from Taiwan with descriptions of three new species Taiwan

Museum Special Publication Series (4) 1ndash25Shen S-C amp Wu K-Y (2011) Cynoglossidae In Shen S-C and Wu K-Y (Eds) Fishes of Taiwan National Museum of

Marine Biology amp Aquarium Pingtung Taiwan pp 763ndash767 (In Chinese)Shinohara G Endo H Matsuura K Machida Y amp Honda H (2001) Annotated Checklist of the deepwater fishes from Tosa

Bay Japan In Fujita T Saito H amp Takeda M (Eds) Deep-sea fauna and pollutants in Tosa Bay National Science Museum Tokyo Monographs Number 20 pp 283ndash343

Shinohara G Sato T Aonuma Y Horikawa H Matsuura K Nakabo T amp Sato K (2005) Annotated checklist of deep-sea fishes from the waters around the Ryukyu Islands Japan In Hasegawa K Shinohara G amp Takeda M (Eds) Deep-sea fauna and pollutants in the Nansei Islands National Science Museum Tokyo Monographs Number 39 pp 683ndash735

Son YH (1980) The fishes of east sea of Chosen Science Press Pyonyang 464 pp (In Korean)Sowerby AdeC (1930) The naturalist in Manchuria Volumes 4ndash5 Tientsin Press 571 ppTamura K Dudley J Nei M amp Kumar S (2007) MEGA4 Molecular Evolutionary Genetics Analysis (MEGA) Software

Version 40 Molecular Biology and Evolution 24(8) 1596ndash1599 httpdxdoiorg101093molbevmsm092Taranetz AJ (1937) Handbook for identification of fishes of Soviet Far East and adjacent waters Bulletin of the Pacific

Science Institute 11 1ndash200 (In Russian)Tautz D Arctander P Minelli A Thomas RH amp Vogler AP (2003) A plea for DNA taxonomy Trends in Ecology and

Evolution 18(2) 70ndash74 httpdxdoiorg101016S0169-5347(02)00041-1Teletchea F (2009) Molecular identification methods of fish species reassessment and possible applications Reviews in Fish

Biology and Fisheries 19(3) 265ndash293 httpdxdoiorg101007s11160-009-9107-4Thompson JD Gibson TJ Plewniak F Jeanmougin F amp Higgins DG (1997) The CLUSTAL_X windows interface

flexible strategies for multiple sequence alignment aided by quality analysis tools Nucleic Acids Research 25(24)

4876ndash4882 httpdxdoiorg101093nar25244876Tian Y Kidokor H amp Watanabe T (2006) Long-term changes in the fish community structure from the Tsushima warm

current region of the JapanEast Sea with an emphasis on the impacts of fishing and climate regime shift over the last four decades Progress in Oceanography 68(2ndash4) 217ndash237 httpdxdoiorg101016jpocean200602009

Tornabene L Baldwin C Weigt LA amp Pezold F (2010) Exploring the diversity of western Atlantic Bathygobius (Teleostei Gobiidae) with cytochrome c oxidase-I with description of two new species Aqua International Journal of Ichthyology 16(4) 141ndash170

Tunnicliffe V Koop BF Tyler J amp So S (2010) Flatfish at seamount hydrothermal vents show strong genetic divergence between volcanic arcs Marine Ecology 31(Suppl 1) 158ndash167 httpdxdoiorg101111j1439-0485201000370x

Victor BC (2007) Coryphopterus kuna a new goby (Perciformes Gobiidae Gobiinae) from the western Caribbean with the identification of the late larval stage and an estimate of the pelagic larval duration Zootaxa 1526 51ndash61

Victor BC (2008) Redescription of Coryphopterus tortugae (Jordan) and a new allied species Coryphopterus bol(Perciformes Gobiidae Gobiinae) from the tropical western Atlantic Ocean Journal of the Ocean Science Foundation2008(1) 1ndash19

Wang C (1993) Ecological characteristics of the fish fauna of the South China Sea In Morton B (Ed) The Marine Biology of the South China Sea Volume 1 Proceedings of the First International Conference on the Marine Biology of Hong Kong and the South China Sea Hong Kong 28 October-3 November pp 77ndash118

Ward RD Zemlak TS Innes BH Last PR amp Hebert PDN (2005) DNA barcoding Australiarsquos fish species Philosophical Transactions of the Royal Society B 360(1462) 1847ndash1857 httpdxdoiorg101098rstb20051716

Ward RD Holmes BH amp Yearsley GK (2008) DNA barcoding reveals a likely second species of Asian seabass (barramundi) (Lates calcarifer) Journal of Fish Biology 72(2) 458ndash463 httpdxdoiorg101111j1095-8649200701703x

Weber M (1913) Die Fische der Siboga-Expedition EJ Brill Leiden 710 ppWeber M amp de Beaufort LF (1929) The fishes of the Indo-Australian Archipelago V Anacanthini Allotriognathi

Heterostomata Berycomorphi Percomorphi families Kuhliidae Apogonidae Plesiopidae Pseudoplesiopidae Priacanthidae Centropomidae The Fishes of the Indo-Australian Archipelago EJ Brill Ltd Leiden Volume 5 458 pp

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Yamada U (2002) Cynoglossidae In Nakabo T (Ed) Fishes of Japan with pictorial keys to the species English edition Tokai University Press Tokyo pp 1388ndash1392

Yasuda F Tokkagi K Tominaga Y Uyeno T Abe T Ishiyama R Iwai T Ochiai A Kuronuma K amp Nakamura M (1981) Dictionary of Japanese fish names and their foreign equivalents Sanseido Company Limited Tokyo Japan 834 pp (In Japanese Chinese English French German Russian and other languages)

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Youn C-H (2002) Fishes of Korea with Pictorial Key and Systematic List Academic Seojeok Seoul 747 pp (In Korean)

Abstract

Symphurus orientalis (Bleeker 1879)

Literature cited

(Eschmeyer 2012) considered a junior subjective synonym of Symphurus Rafinesque 1810 and this generic assignment is followed in this study] Information provided in the original description and derived from the accompanying illustration of S orientalis distinguishes this species as a tonguefish featuring 100 dorsal-fin rays 86 anal-fin rays 12 caudal-fin rays approximately 90 longitudinal scales and 40 transverse scales and with an ocular-side pigmentation pattern consisting of multiple crossbands a uniformly white blind side and black peritoneum

Somewhere in its history the holotype of S orientalis was lost and its whereabouts are unknown (Eschmeyer 2012) Limited diagnostic information contained in the original description compounded by loss of this holotype have rendered it difficult for subsequent researchers interested in the taxonomy and systematics of Indo-West Pacific tonguefishes to adequately distinguish the species described by Bleeker (1879) Of characters discussed in the original description noteworthy is the number of caudal-fin rays (12) reported for the holotype as this is one of the most important characters for differentiating phenetic groups of species within Symphurus (Ginsburg 1951 Chabanaud 1955 Munroe 1992) In an extensive study summarizing information from several thousand specimens of Symphurus Munroe (1992) reported that the number of caudal-fin rays is one of the most conservative and important diagnostic features for identifying species of Symphurus

Several other internal morphological characteristics of symphurine tonguefishes also important in characterizing and diagnosing these species include the pattern of interdigitation of dorsal proximal pterygiophores and neural spines (ID pattern Munroe 1992) and the number of abdominal and total vertebrae (Chabanaud 1955 Munroe 1992) Indo-Pacific species of Symphurus can readily be separated into different phenetic groups based on combinations of the numbers of caudal-fin rays abdominal vertebrae and by their ID patterns (Munroe 1992) Internal features were not reported in the original description of A orientalis and it is not now possible to know what the character states were for this nominal species Thus the species concept of S orientalis as envisioned by Bleeker (1879) is known only from the limited information available in the original description and the accompanying illustration of this nominal species

Reliance on data presented in the original description of S orientalis and the inability to compare the holotype specimen with other nominal species of Symphurus collected subsequently has resulted in considerable taxonomic confusion and misidentifications regarding specimens of Symphurus collected in the Indo-Pacific region Jordan and Starks (1906) for example in their review of the flounders and soles of Japan questioned whether the specimen described by Bleeker actually belonged to Symphurus Okada and Matsubara (1938) and Matsubara (1955) wrote identification keys for fishes occurring in Japanese waters but did not list any catalogue information for specimens they examined In their key to the species of Symphurus they provided ranges for meristic and morphometric features for specimens purportedly of S orientalis However based on specimens we identified as S orientalis (Lee amp Munroe unpubl data) data in Matsubararsquos study overestimates those for S orientalis and appear to be derived from at least two and possibly more sympatric and morphologically similar species of Symphurus featuring 12 caudal-fin rays including some specimens with meristic features much lower than those of S orientalis (Bleeker 1879 Lee amp Munroe unpubl data)

The works of Okada and Matsubara (1938) and Matsubara (1955) had a profound influence on another work that of Ochiai (1959 Japanese edition and English edition 1963) which purportedly redescribed S orientalis Interestingly Ochiairsquos redescription which is based on catalogued specimens reports ranges of several important meristic and morphometric features for this species that are the same as those listed in Matsubararsquos study (1955) It is uncertain if Ochiairsquos work was based on the same specimens as those included in Matsubararsquos study (1955) because catalogue numbers for specimens were not included in the latter study but it seems likely Ochiairsquos study (1959 1963) is the most widely cited work on S orientalis but his redescription includes data from several morphologically similar species with 12 caudal-fin rays A consequence of being so widely cited is that the erroneous data in Ochiairsquos study has been perpetuated in subsequent literature based on this work and this has contributed significantly to the confusion and misunderstanding regarding the range of variation in morphological features in this nominal species

Chen and Weng (1965) based on four specimens potentially represents the first report of S orientalis from Taiwanese waters but according to the number of caudal-fin rays (15) they reported for their specimens it is uncertain whether they actually examined any specimens of this species Specimens included in the study by Chen and Weng (1965) have been lost so it is difficult to know whether these authors counted caudal-fin rays accurately or if their redescription was based on specimens of more than one species at least one of which featured 15 caudal-

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

Literature cited

ID Pattern

1-2-2-2-2 1-2-3-2-2 1-2-2-2-1 1-3-2-2-2 1-2-1-2-2 N

83 3 1 2 2 91

Dorsal-fin rays

96 97 98 99 100 101 N

12 11 23 23 11 12 92

Anal-fin rays

82 83 84 85 86 87 88 89 N

2 8 19 15 26 17 4 1 92

11 12 N 3+6 N

2 92 94 93 93

Total vertebrae

52 53 54 55 N

5 27 50 9 91

87 88 89 90 91 92 93 94 95 96 97 98 99 N

1 4 2 6 6 9 9 13 7 10 7 9 3 86

Head scale count

18 19 20 21 22 N

5 31 33 16 1 86

37 38 39 40 41 42 N

7 21 21 24 12 1 86

Discussion

Acknowledgments

Literature cited

Abstract

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Discussion

Acknowledgments

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Abstract

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Discussion

Acknowledgments

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Abstract

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Discussion

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Discussion

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Discussion

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Discussion

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Discussion

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Discussion

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Documents

Symphurus orientalis (Bleeker) redefined based on ...€¦ · ISSN 1175-5326 (print edition) ... Symphurus orientalis (Bleeker) redefined based on morphological and molecular characters