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Eptatretus strickrotti n. sp. (Myxinidae): first hagfish captured from a hydrothermal vent.

Introduction

The number of recognized genera in the subfamily Eptatretinae is still controversial, with various specialists recognizing only Eptatretus Cloquet, 1819 (e.g., Strahan, 1975; Fernholm, 1998; Chen et al., 2005), Eptatretus and Paramyxine Dean, 1904 (e.g., Mok, 2001), Eptatretus and Quadratus Wisner, 1999 (e.g., Froese and Pauly, 2005), or Eptatretus, Paramyxine, and Quadratus (e.g., Wisner, 1999; McMillan and Wisner, 2004; Mok and McMillan, 2004). This lack of a consensus seems to be caused by the paucity of phylogenetic studies, which are surprisingly few (Jansson et al., 1995; Kuo et al., 2003; Chen et al., 2005) given the importance of jawless fishes in vertebrate evolution, medical research, and fisheries (see, e.g., Nelson, 2006; Ota and Kuratani, 2006). The subfamily Eptatretinae has been extensively revised within the last 25 years (e.g., Fernholm and Hubbs, 1981; Kuo et al., 1994; McMillan and Wisner, 2004)--almost tripling the number of known species from about 16 in 1980 to 46 currently (including the one herein described). The reasons for this increase are many, but a significant number of new species have been discovered after capture from submersibles in remote rocky habitats where traditional fishing methods are less successful (Mincarone and McCosker, 2004). So far, however, submarine investigations of hydrothermal vents have not captured any myxinids from these unique environments (Biscoito et al., 2002; Wolf, 2005; Desbruyeres et al., 2006), and only a single unconfirmed record exists in the literature (E. okinoseanus from Okinawa Trough [Ohta and Kim, 2001]).

During the Easter Microplate expedition, 12 March-6 April 2005, with R/V Atlantis and DSV Alvin, a single myxinid specimen was caught by the Alvin slurp gun at the East Pacific Rise vent site known as 38[degrees]S (a video clip of the capture is available upon request from the authors). The specimen does not match any known species of hagfish and is here described as a new species of Eptatretus. Its phylogenetic position was determined using Bayesian analysis of molecular 16S rRNA data.

Materials and Methods

Methods of counting and anatomical terminology follow Fernholm and Hubbs (1981) and McMillan and Wisner (1984, 2004). Body width is measured at the pharyngocutaneous duct (PCD) following McMillan and Wisner (2004), and not McMillan and Wisner (1984) in which body width is the maximum width between the rostrum and PCD. Body depth is measured at the PCD and therefore does not follow McMillan and Wisner (2004), in which depth is maximum body depth. The reason for this choice is to maintain consistency in the measurement of body depth; however, for the present species, depth at the PCD and maximum depth are similar. Terminology of nasal sinus papillae follows Mok (2001), except for the lateral structures in the nasal sinus floor, which are here referred to simply as papillae, not longitudinal folds. Since only one specimen is available for examination, no attempt was made to dissect the nasal sinus as suggested by Mok (2001); we made only direct observations through the nasal opening.

Author names of species are provided in the Appendix. "Subgenera" abbreviations Eptatretus E., Eptatretus P., and Eptatretus Q. are used to illustrate that some authors recognize the genera Paramyxine and Quadratus and to facilitate comparison with earlier literature that uses these genera. Comparative material includes Eptatretus E. deani, KU 28249 (Southern California, 34[degrees]47'N, 120[degrees]85'W); E. E. longipinnis, SAMA F4042 Holotype, SAMA F7540 (South Australia, Port MacDonnell, ca. 38[degrees]S, 141[degrees]E), SAMA F5676, SAMA F3611; E. E. cirrhatus, ZMUC P02221-228 (Milford Sound, New Zealand). Institutional abbreviations: CAS = California Academy of Sciences; KU = University of Kansas; SAMA = South Australian Museum; ZMUC = Zoological Museum, University of Copenhagen.

Molecular sequencing

Phylogenetic analyses were conducted with a combination of published and original DNA sequences. We list the origin of specimens (geographical coordinates) used to obtain original sequences in Table 1 along with their voucher (museum) and GenBank accession numbers. To obtain the original sequences, we extracted genomic DNA from ethanol-preserved muscle tissue (50 mg) that was treated with the Qiagen Dneasy isolation kit, according to manufacturer's instructions (Qiagen Inc., Valencia, CA). Polymerase chain reaction (PCR) conditions for amplification of 16S rRNA contained 30-100 ng of template DNA, 5 [micro]l of 10X buffer (supplied by manufacturer), 5 [micro]l of Mg[Cl.sub.2] (2.5 [micro]mol [1.sup.-1]), 2 [micro]l of each primer (10 [micro]m final conc.), 2.5 units of Taq polymerase (Promega Inc., WI), 5 [micro]l of a 2 mmol [1.sup.-1] stock solution of dNTPs, and sterile [H.sub.2]O to a final volume of 25 [micro]l. PCR was performed with a Cetus 9600 DNA thermocycler (Perkin-Elmer Corporation, Connecticut) using the conditions of Kuo et al. (2003).

PCR products were purified using a Qiagen PCR purification kit (Qiagen Inc., Valencia, CA) or Montage columns (Millipore, Billerica, MA). The purified template DNA was sequenced using a Big Dye Terminator cycle sequencing reaction kit (PE Biosystems, Foster City, CA) and ABI Prism 3100 DNA sequencers (Applied Biosystems Inc., Foster, CA). PCR products were sequenced bidirectionally from each individual sample, using the same forward and reverse primers as used in PCR. DNA sequence alignments were initially constructed using Sequencher (Gene Codes Corp. Inc., Ann Arbor, MI). Secondary structure of rRNA (i.e., stems and loops) was inferred using the program GeneBee (Brodsky et al., 1992). The 16S rRNA alignment analyzed in this study is available in GenBank PopSet (accession numbers in Table 1).

Phylogenetic analyses

Bayesian phylogenetic trees were estimated for 16S rRNA using MrBayes version 3.0b4 (Huelsenbeck and Ronquist, 2001), partitioning by structure (stem vs. loop). Representatives of Myxine (Table 1) and Petromyzon marinus were chosen as outgroup taxa based on Kuo et al. (2003). Parameters are available upon request (WJJ). The Monte Carlo Markov chain (MCMC) length was 1.1 X [10.sup.6] generations with 6 markov chains, and we sampled the chain every 100 generations to minimize autocorrelation. MCMC convergence was assessed by visually inspecting the sample paths of model parameters (to determine an appropriate burn-in period) and by repeating the analysis multiple times (at least 3) with random initial parameter values (to assess the dependence of posterior distributions on initial conditions).

Parameter estimates were graphically analyzed to assess stability. Log-likelihood values and associated parameters for sampled trees stabilized after approximately 5-6 X [10.sup.5] generations. Therefore, we conservatively used the last 5000 sampled trees to estimate Bayesian posterior probabilities (BPP). If [greater than or equal to]95% of the sampled trees contained a given clade, we considered it to be significantly supported by our data (sensu Wilcox et al., 2002).

Eptatretus strickrotti new species

Figures 1-4

Holotype

CAS 223480, 314 mm total length (TL), sex unknown, East Pacific Rise, 37[degrees]47.363'S, 110[degrees]54.905'W, 2211 m, 21:24 GMT, R/V Atlantis, DSV Alvin dive 4089 (38[degrees]S), 23 March 2005, Starboard observer: Daniel Layton-Matthews, Port observer: William Jones, Pilot: Bruce Strickrott. Specimen caught, using the slurp gun/suction sampler on the Alvin, while swimming about 1 m above the bottom of lobate basalt flow along a fissure near Sebastian's Steamer hydrothermal vent (Fig. 1A).

Diagnosis

Distinguished from all congeners by the slender body, depth at PCD 2.9% TL, paired and median ventral nasal sinus papillae, and presence of 10 afferent branchial arteries on the medial ventral aorta. It is further characterized by the combination of the following characters: slime pores--prebranchial 18, branchial 12, trunk 70, caudal 19, total 119; gill pouches (GP) and gill apertures (GA) 12, PCD widely separated from the opening of posterior left GAs; multicusps in anterior row 3 and multicusps in posterior row 2, total cusps 46; paired dorsal nasal sinus papillae; eyespot absent; body coloration pink.

Description

Body very slender, laterally compressed, and oval in cross section. Width/depth ratio at PCD 0.6. Rostrum rather long, rounded (Fig. 2A, B). Two bilaterally symmetrical, bi-tipped, crescent-shaped papillae present in the dorsal surface of the nasal tube (nasal sinus). In the floor of the nasal tube, a large, median, pillow-shaped papilla is present in addition to two bilaterally symmetrical, single-tipped papillae (Fig. 3). Eyespots and head grooves absent. Ventral finfold vestigial, maximum depth about 0.8 mm, beginning approximately at the middle of the body and extending posteriorly to the cloaca. Caudal finfold thin and low (missing about the last 3 mm), extending around the tail to dorsal surface, ending nearly above the cloaca. Tentacles relatively large, the first and third almost equal in length (1.4% and 1.5% TL, respectively), the second a bit shorter (1.1% TL) (Fig. 2A, B).

Dental muscle pale and short: length 17.8% TL, width 1.5% TL, only slightly overlapping with anteriormost gill pouch, resulting in the two rows of GPs lying closely together (Fig. 2E).

Twelve GPs and GAs, the posterior left GA widely separated from the opening of the PCD. The distance (1.4 mm) is about the same as that between the last two GAs (Fig. 2D). All efferent branchial ducts of about similar length. Ventral aorta (VA) bifurcating at an anterior position between 2nd and 3rd GP (counted from the snout toward the heart). Ten afferent branchial arteries on each side of the VA posterior to bifurcation. Each separated ventral aorta (SVA) has one afferent branchial artery and ends in the anteriormost (1st) GP.

Palatine cusp short (length 0.94 mm, width at base 0.44 mm). First 3 cusps in anterior row and first 2 cusps in posterior row fused at their bases (= multicusp configuration 3/2), followed by 9 unicusps in both rows. Total cusps 46. Longest cusp in anterior row is first unicusp next to multicusps, 1.7 mm long, 0.7 mm at base; longest cusp in posterior row is first unicusp next to multicusps, 1.5 mm long, 0.4 mm at base.

Slime pores very small and difficult to count, especially in trunk region. Prebranchial slime pores 18, branchial slime pores 12, trunk slime pores 74, caudal slime pores 19 (2 of which are above the cloaca).

Coloration. Live specimen bright pink, with a narrow, darker red stripe along the dorsal surface (Fig. 1A). Preserved color pale, light yellow, with some internal dark pigmentation seen through skin, in the head and along mid-body (Fig. 1B). Specimens seen from submersible pale, whitish, probably due to the light source of the submersible.

Comparisons. Selected characters are provided for eptatretines in the Appendix. Eptatretus E. strickrotti is a very distinct species of hagfish, differing from all congeners by the very slender body (depth at PCD = max. depth 2.9% vs. 4.0%-15.3% TL) and numerous afferent branchial arteries from medial part of the VA (10 vs. 0-6) (Mok and McMillan, 2004).

It differs from the species often assigned to Paramyxine (8 species) and Quadratus (4 species) by having a high number of GPs and GAs (12 vs. 5-6 and 5-6), in a straight line (vs. fairly straight and nonlinear, crowded), corresponding long branchial region (9.3% vs. 1.3%-5.6% and 1.1%-2.8% TL), and in the number of caudal slime pores (19 vs. 5-14 and 7-12) (Appendix).

Eptatretus E. strickrotti resembles eight East Pacific congeners (E. E. bischoffli, E. E. deani, E. E. fritzi, E. E. mcconnaugheyi, E. E. nanii, E. E. polytrema, E. E. sinus, and E. E. stoutii) (see Appendix) by having a high (12 and 9-14) number of GPs and GAs and a long branchial region (9.3% and 9.2%-22.0% TL), but it differs from all these by the number of slime pores (e.g., caudal 19 vs. 7-17 and total 119 vs. 66-88), presence of dorsal (and ventral) nasal sinus papillae (vs. absent), lack of visible eye spot (vs. present), length of dental muscle (18% vs. 20%-39% TL), and anteritorly bifurcated VA (at GP 2-3 vs. 4-13) (Appendix). It is further separated from E. E. bischoffii, E. E. nanii, and E. E. polytrema from off Chile by the number of multicusps in the posterior row (2 vs. 3) and several body proportions (e.g., trunk length 59% vs. 45%-54% TL). Five Northeast American species (E. E. deani, E. E. fritzi, E. E. mcconnaugheyi, E. E. sinus, and E. E. stoutii) have 3/2 multicusps like E. E. strickrotti. In addition to the above-mentioned characters, however, E. E. strickrotti differs from these five in the body color (pink vs. gray, brown, or black, except for a few specimens of E. E. stoutii with pinkish overtones [Wisner and McMillan, 19901 and a single pink E. E. deani [McMillan, 1999]).

Eptatretus E. strickrotti resembles 11 of the remaining 24 species of Eptatretus by the 3/2 multicusp configuration, but it differs from all these by the more numerous gill pouches (12 vs. 5-8) and in total number of slime pores (119 vs. 72-110 or 128-130) (Appendix). Most similar slime-pore counts are seen in three Pacific deep-water species: E. E. carlhubbsi from off Hawaii, Guam and Wake Islands; E. E. laurahubbsae from off Chile; and E. E. eos Fernholm, 1991, from the Tasman Sea (e.g., trunk slime pores 70, 60-70, 60-67, and 70-75, respectively). Eptatretus E. strickrotti further resembles E. E. carlhubbsi and E. E. laurahubbsae in the paired dorsal sinus papillae, but differs in multicusp configuration (3/2 vs. 2/3 and 2/2) and by a much lower number of unicusps (46 vs. 64-71 and 61-68). Eptatretus E. strickrotti also resembles E. E. eos in the pink color, but differs in the number of slime pores (e.g., caudal pores 19 vs. 26-27), in addition to the aforementioned characters.

Etymology. Named in honor of DSV Alvin pilot Mr. Bruce Strickrott, in recognition of his expertise with Alvin and slurp-gun capture of the holotype specimen and other mobile hydrothermal vent animals.

Distribution. Known from the holotype only, caught at the East Pacific Rise, vent site (38[degrees]S), at 2211-m depth. This is the first member of the jawless fishes to be captured from a hydrothermal vent site, confirming that they do occur in this type of habitat. Hagfishes are often difficult to identify from photographs and are easily confused with synaphobranchid eels and even wormlike invertebrates. Ohta and Kim (2001) reported observations of specimens of Eptatretus E. okinoseanus from a vent on Iheya Ridge, Okinawa Trough, but neither illustrations nor text justifying the identification were provided. Eptatretus E. strickrotti is the fourth species of Eptatretus recorded from depths below 2000 m: E. E. deani (2743 m), E. E. fritzi (2743 m), and E. E. laurahubbsae (2400 m) (Fernholm, 1998).

During the French Biospeedo expedition (Jollivet et al., 2004) to the East Pacific Rise in 2004, at least seven specimens of myxinids were observed and filmed on the vent sites Grommit (21[degrees]33.664'S, 114[degrees]17.982'W, 2838 m), Oasis (17[degrees]25.38'S, 113[degrees]12.29'W, 2586 m), and Yaquina (07[degrees]29.96'S, 107[degrees]53.95'W, 2700 m), thus extending the range for East Pacific Rise myxinids 30[degrees] to the north of the holotype catch site. Those observations also extended the depth range by about 800 m. Since none of these specimens were collected, we cannot be sure if they belong to the species here described. They have a similar slender body and appear pale to white on the video recordings, which is not in conflict with the light pink color of the collected specimen. Two myxinids were seen swimming toward the current like the captured specimen, and two were drifting with the current together with unidentified ophidiiform species.

In general, the obligate vent fish fauna of the East Pacific Rise is very poorly known, with only about 30 specimens of Thermaces spp. collected (Geistdoerfer and Seuront, 1995) and two specimens each of Thermichthys hollisi (Nielsen and Cohen, 2005) and Ventichthys biospeedoi (Nielsen et al., 2006).

Phylogenetic analysis

The complete aligned dataset consisted of 571 base pairs: 315 positions were designated as stems and 256 as loops. Three new 16S rDNA sequences were obtained in the present study: 267 sites were variable, including 123 that were parsimony informative.

The Bayesian analysis generated a topology in the monophyletic Eptatretus clade that was in accordance with the parsimony and neighbor-joining trees presented by Kuo et al. (2003) and Chen et al. (2005) (Fig. 4). The 16S rRNA phylogeny supports placement of Eptatretus in a reciprocally monophyletic clade relative to Myxine (Fig. 4). Resolution within Eptatretus was generally poor, as observed in Kuo et al. (2003) and Chen et al. (2005). Eptatretus P. cheni was resolved as a basal node but not highly supported (BPP = 0.63). Placement of E. E. strickrotti n: sp. as the most basal member of the Eptatretus clade was supported, but not strongly. Additional new 16S rRNA sequences collected in the present study include E. E. deani, which is strongly supported as the sister species to E. E. stoutii. Finally, E. E. longipinnis is placed with E. Q. yangi and E. P. sheni, though the support for these respective nodes are weak (Fig. 4).

Discussion

The gill pouch-gill aperture relationship has been extensively studied and has been used to establish genera and even subfamilies (Wisner, 1999). The present results with Eptatretus E. strickrotti in the most basal position add support to the view that a simple condition with the anterior gill aperture almost beside the anterior gill pouch is the plesiomorphic state, whereas a posterior origin of the gill apertures represents the apomorphic state. Other basal taxa with the simple condition are E. E. cirrhatus, E. E. stoutii, and E. E. deani, whereas the basal E. cheni contradicts this pattern by having a posterior origin of the first gill aperture, typical for the terminal taxa sometimes referred to as Paramyxine or Quadratus. In the present analysis, none of these form monophyletic groups, and we therefore support the synonymization of these genera in Eptatretus, as suggested by Fernholm (1998), Kuo et al. (2003), and Chen et al. (2005).

In a few myxinids, the PCD is not confluent with the last gill aperture. The present results with E. E. strickrotti and E. P. cheni in basal positions suggest that a separate PCD and last gill aperture is likely to be plesiomorphic in eptatretines, as it is expected to have been in the common ancestor of all hagfishes. More material is needed to show whether the condition is stable as in Notomyxine tridentiger (Garman, 1899), E. E. bischoffli, E. P. cheni, and E. P. fernholmi or variable as in several other species (e.g., E. E. burgeri, E. E. nanii, and E. E. polytrema) (Wisner and McMillan, 1988; McMillan and Wisner, 2004).

Branching of the ventral aorta (VA) is considered synapomorphic for Eptatretinae (Mok and McMillan, 2004), in contrast to the plesiomorphic unbranched condition in Myxininae. Chen et al. (2005) argued that the transformation series suggested by Mok and McMillan (2004) (i.e., bifurcation from a site midway between the heart and the 1st gill pouch [e.g., E. E. burgeri] toward a site close to the heart [e.g., E. E. chinensis, E. E. cirrhatus, E. P. sheni]) was not in conflict with their molecular data regarding E. E. chinensis and E. P. sheni. However, they did not comment on the fact that E. E. cirrhatus, which has a bifurcation very close to the heart, was placed in a basal position in their phylogeny. In the present analysis, the basal E. E. strickrotti with VA bifurcation close to the anterior gill pouches indicates that this is the plesiomorphic condition for eptatretines. We propose that to express this character by the GP number where the VA bifurcates gives a poor polarization of the character because of the large variation in the total number of GPs. We therefore suggest that it be expressed as the number of the GP at which bifurcation occurs divided by the total number of GPs (Appendix).

The dorsal nasal sinus papillae have proved to be specific and useful for resolving taxonomic problems in Myxinidae (Mok, 2001; Moller et al., 2005). The present phylogenetic reconstruction confirms that presence of paired dorsal papillae is the plesiomorphic state in Eptatretinae. They are present in E. E. strickrotti, E. P. cheni, and E. E. cirrhatus, and absent in the other, more terminal Eptatretus species surveyed. This is in agreement with Mok (2001), who suggested that the common ancestor of hagfishes could have had these papillae. With only one specimen available, it was not possible for us to check whether the papillae in E. E. strickrotti have the plesiomorphic supporting cartilage as in Myxininae and E. P. cheni.

Acknowledgments

The expedition during which the new hagfish was caught was made possible by National Science Foundation grants to Dr. Robert Vrijenhoek (NSF OCE-0241613) and Dr. Cindy Van Dover (NSF OCE-0350554). We thank Karen Jacobsen for making the drawing; Dr. Didier Jollivet and Dr. Michel Segonzac (IFREMER) for providing video recordings from the Biospeedo expedition; Dr. Andy Bentley and Dr. Ed Wiley (Natural History Museum & Biodiversity Research Center, Kansas) for donating a tissue sample of E. E. deani; and Dr. Terry Bertozzi, Dr. Ralf Foster, and Dr. Steve Donnellan (South Australian Museum) for sharing extracted DNA of E. E. longipinnis.

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Wilcox, T. P., D. J. Zwickl, T. A. Heath, and D. M. Hillis. 2002. Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Mol. Phylogenet. Evol. 25: 361-371.

Wisner, R. L. 1999. Descriptions of two new subfamilies and a new genus of hagfishes (Cyclostomata: Myxinidae). Zool. Stud. 38: 307-313.

Wisner, R. L., and C. B. McMillan. 1988. A new species of hagfish, genus Eptatretus (Cyclostomata, Myxinidae), from the Pacific Ocean near Valparaiso, Chile, with new data on E. bischoffii and E. polytrema. Trans. S. Diego Soc. Nat. Hist. 21: 227-244.

Wisner, R. L., and C. B. McMillan. 1990. Three new species of hagfishes, genus Eptatretus (Cyclostomata, Myxinidae), from the Pacific Coast of North America, with new data on E. deani and E. stoutii. Fish. Bull. 88: 787-804.

Wolf, T. 2005. Composition and endemism of the deep-sea hydrothermal vent fauna. Cah. Biol. Mar. 46: 97-104.

Wongratana, T. 1983. Eptatretus indrambaryai, a new species of hagfish (Myxinidae) from the Andaman Sea. Nat. Hist. Bull. Siam Soc. 31: 139-150.

Appendix
Appendix Table 1. Selected characters of Eptatretus spp.

                                   Gill     Fused  Total  Total slime
Species                            pouches  cusps  cusps  pores

E. Q. ancon Mok. Saavedra-Diaz &    6       3/2    60      80
  Acero-P, 2001
E. P. atami (Dean, 1904)            6       3/3    47-52   71-78
E. E. burgeri# (Girard, 1855)       5-6     3/2    35-42   81-92
E. P. cheni# (Shen and Tao, 1975)   5       3/3    50-53   75-81
E. E. bischoffli (Schneider,       10-11    3/3    44-52   74-83
  1880)
E. E. caribbeaus Fernholm, 1982     7       3/3    54-58   79-85
E. E. carlhbbsi McMillan and        7       2/3    64-71   93-110
  Wisner, 1984
E. E. chinensis# Kuo and Mok,       6       3/3    46-52   72-83
  1994
E. E. cirrhatus# (Forster, 1801)    7       3/3    43-51   79-90
E. E. deani# (Evermann and         10-12    3/2    37-46   67-80
  Goldsborough, 1907)
E. E. eos Fernholm, 1991            5       3/2-3  34     128-130
E. E. fernholmi McMillan and        8       3/2    51      81
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and      6       3/2    38-44   64-71
  Mok, 1994)
E. E. fritzi Wisner and            10-12    3/2    38-46   74-85
  McMillan, 1990
E. E. goliath Mincarone and         7       3/3    54      92
  Stewart, 2006
E. E. grouseri McMillan, 1999       5-6     3/2    44-48   71-79
E. E. hexatrema (Muller, 1836)      5-6     3/2    40-48   90-105
E. E. indrambaryai Wongratana,      8       3/2    45-48   77-82
  1983
E. E. lakeside Mincarone and        5       3/3    36      88
  McCosker, 2004
E. E. laurahubbsae McMillan and     7       2-3/2  61-68   97-105
  Wisner, 1984
E. E. longipinnis# Strahan, 1975    6       3/2    30     104-108
E. E. mcconnaugheyi Wisner and     12-14    3/2    42      67-84
  McMillan, 1990
E. E. mccoskeri McMillan, 1999      8       3/3    48-51   72-74
E. E. mendozai Hensley, 1985        6       3/3    56-61   77-82
E. E. menezesi Mincarone, 2000      7       3/3    52-60   86-94
E. E. minor Fernholm and Hubbs,     5-6     3/3    46-54   74-82
  1981
E. P. moki (McMillan and Wisner,    6       3/2    38-42   75-82
  2004)
E. E. mulitidens Fernholm and       6       3/3    52-57   87-91
  Hubbs, 1981
E. E. nanii Wisner and McMillan,   12-13    3/3    49-55   72-82
  1988
E. Q. nelsoni# (Kuo, Huang and      3-5     3/2    32-40   57-67
  Mok, 1994)
E. E. octatrema (Barnard, 1923)     8       3/2    40     102-103
E. E. okinoseanus (Dean, 1904)      8       3/2    40-49   87-97
E. E. polytrema (Girard, 1855)     13-14    3/3    45-51   72-79
E. E. profundus (Barnard, 1923)     5       3/2    42      82-83
E. P. sheni# (Kuo, Huang and        6       3/3    48-53   62-74
  Mok, 1994)
E. E. sinus Wisner and McMillan,    9-12    3/2    34-46   66-82
  1990
E. P. springeri (Bigelow and        6       3/2    48-52   84-92
  Schroeder, 1952)
E. E. stoutii# (Lockington, 1878)  10-14    3/2    36-46   71-88
E. E. strahani McMillan and         7       3/3    47-52   76-80
  Wisner, 1984
E. E. strickrotti# n. sp.          12       3/2    46     119
E. Q. taiwanae# (Shen and Tao,      6       3/2    32-40   60-68
  1975)
E. P. walkeri (McMillan and         5-6     3/2    38-44   69-79
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz      5       3/2    41-43   73-75
  and Acero, 2001
E. E. wisneri McMillan, 1999        8       3/2    44      76
E. P. wisneri (Kuo, Hang and        6       3/2    36-44   63-72
  Mok, 1994)
E. Q. yangi# (Teng, 1958)           5       3/2    32-40   68-79

                                           GP at
                                   Dorsal  end    ABA at  VA bifurcates
Species                            NSP     of DM  MVA     at GP

E. Q. ancon Mok. Saavedra-Diaz &   --      --      3       4-5
  Acero-P, 2001
E. P. atami (Dean, 1904)           absent  1-2     2       5-6
E. E. burgeri# (Girard, 1855)      absent  1-2     3       3-4
E. P. cheni# (Shen and Tao, 1975)  paired  1-2     3       3
E. E. bischoffli (Schneider,       absent  6-11    0      10-11
  1880)
E. E. caribbeaus Fernholm, 1982    absent  --      0      --
E. E. carlhbbsi McMillan and       paired  2-4     2       7
  Wisner, 1984
E. E. chinensis# Kuo and Mok,      absent  2-3     0-1     5-6
  1994
E. E. cirrhatus# (Forster, 1801)   paired  4-7     0       7
E. E. deani# (Evermann and         absent  5       5       6
  Goldsborough, 1907)
E. E. eos Fernholm, 1991           --      --     --      --
E. E. fernholmi McMillan and       --      --     --      --
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and     absent  1-3     3       3-4
  Mok, 1994)
E. E. fritzi Wisner and            absent  5       3       9
  McMillan, 1990
E. E. goliath Mincarone and        paired  4       0       7
  Stewart, 2006
E. E. grouseri McMillan, 1999      --      1-2    --       5
E. E. hexatrema (Muller, 1836)     paired  --      3      --
E. E. indrambaryai Wongratana,     --      2-3     0       8
  1983
E. E. lakeside Mincarone and       paired  0       3       2
  McCosker, 2004
E. E. laurahubbsae McMillan and    paired  2-5     3       5-7
  Wisner, 1984
E. E. longipinnis# Strahan, 1975   absent  2-3     2-3     4
E. E. mcconnaugheyi Wisner and     absent  9       1-3    11
  McMillan, 1990
E. E. mccoskeri McMillan, 1999     --      4-6     2       6-7
E. E. mendozai Hensley, 1985       paired  2-4     0       5-6
E. E. menezesi Mincarone, 2000     --      2-5     2       5-6
E. E. minor Fernholm and Hubbs,    paired  3-4     0-1     5-6
  1981
E. P. moki (McMillan and Wisner,   --      1-2    --       3-4
  2004)
E. E. mulitidens Fernholm and      paired  2-3     0       6
  Hubbs, 1981
E. E. nanii Wisner and McMillan,   absent  4-7     6       4-9
  1988
E. Q. nelsoni# (Kuo, Huang and     absent  1-2     2       3-4
  Mok, 1994)
E. E. octatrema (Barnard, 1923)    --      --     --      --
E. E. okinoseanus (Dean, 1904)     absent  1-3     2       7-8
E. E. polytrema (Girard, 1855)     absent  5-10    1       9-13
E. E. profundus (Barnard, 1923)    absent  --     --       2-3
E. P. sheni# (Kuo, Huang and       absent  3-4     0-1     4-5
  Mok, 1994)
E. E. sinus Wisner and McMillan,   absent  2       6       5-6
  1990
E. P. springeri (Bigelow and       --      1-3     1       4-5
  Schroeder, 1952)
E. E. stoutii# (Lockington, 1878)  absent  3       4       7
E. E. strahani McMillan and        paired  3-5     0       7
  Wisner, 1984
E. E. strickrotti# n. sp.          paired  1      10       2-3
E. Q. taiwanae# (Shen and Tao,     absent  1-2     3       4-5
  1975)
E. P. walkeri (McMillan and        --      1-2    --       3-4
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz     --      0       3-5     1-2
  and Acero, 2001
E. E. wisneri McMillan, 1999       --      3      --       5
E. P. wisneri (Kuo, Hang and       absent  2-4     3       3-4
  Mok, 1994)
E. Q. yangi# (Teng, 1958)          absent  1-2     2-3     4-5

                                   VA bifurcates at
Species                            GP/total GP's     GA configuration

E. Q. ancon Mok. Saavedra-Diaz &   0.7-0.8           crowded
  Acero-P, 2001
E. P. atami (Dean, 1904)           0.8-1.0           fairly straight
E. E. burgeri# (Girard, 1855)      0.6-0.7           linear
E. P. cheni# (Shen and Tao, 1975)  0.6               fairly straight
E. E. bischoffli (Schneider,       1.00              linear
  1880)
E. E. caribbeaus Fernholm, 1982    --                linear
E. E. carlhbbsi McMillan and       1.0               linear
  Wisner, 1984
E. E. chinensis# Kuo and Mok,      0.8-1.0           linear
  1994
E. E. cirrhatus# (Forster, 1801)   1.0               linear
E. E. deani# (Evermann and         0.5-0.6           linear
  Goldsborough, 1907)
E. E. eos Fernholm, 1991           --                linear
E. E. fernholmi McMillan and       --                linear
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and     0.5-0.7           fairly straight
  Mok, 1994)
E. E. fritzi Wisner and            0.8-0.9           linear
  McMillan, 1990
E. E. goliath Mincarone and        1.0               linear
  Stewart, 2006
E. E. grouseri McMillan, 1999      0.8-1.0           linear
E. E. hexatrema (Muller, 1836)     --                linear
E. E. indrambaryai Wongratana,     1.0               linear
  1983
E. E. lakeside Mincarone and       0.4               linear
  McCosker, 2004
E. E. laurahubbsae McMillan and    0.7-1.0           linear
  Wisner, 1984
E. E. longipinnis# Strahan, 1975   0.7               linear
E. E. mcconnaugheyi Wisner and     0.8               linear
  McMillan, 1990
E. E. mccoskeri McMillan, 1999     0.8-0.9           linear
E. E. mendozai Hensley, 1985       0.8-1.0           linear
E. E. menezesi Mincarone, 2000     0.7-0.9           linear
E. E. minor Fernholm and Hubbs,    0.8-1.0           linear
  1981
E. P. moki (McMillan and Wisner,   0.5-0.7           fairly straight
  2004)
E. E. mulitidens Fernholm and      1.0               linear
  Hubbs, 1981
E. E. nanii Wisner and McMillan,   0.3-0.7           linear
  1988
E. Q. nelsoni# (Kuo, Huang and     0.6-0.8           crowded
  Mok, 1994)
E. E. octatrema (Barnard, 1923)    --                linear
E. E. okinoseanus (Dean, 1904)     0.9-1.0           linear
E. E. polytrema (Girard, 1855)     0.7-0.9           linear
E. E. profundus (Barnard, 1923)    0.4-0.6           linear
E. P. sheni# (Kuo, Huang and       0.7-0.8           fairly straight
  Mok, 1994)
E. E. sinus Wisner and McMillan,   0.5-0.6           linear
  1990
E. P. springeri (Bigelow and       0.7-0.8           fairly straight
  Schroeder, 1952)
E. E. stoutii# (Lockington, 1878)  0.5-0.7           linear
E. E. strahani McMillan and        1.0               linear
  Wisner, 1984
E. E. strickrotti# n. sp.          0.2-0.23          linear
E. Q. taiwanae# (Shen and Tao,     0.7-0.8           crowded
  1975)
E. P. walkeri (McMillan and        0.5-0.7           fairly straight
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz     0.2-0.4           linear
  and Acero, 2001
E. E. wisneri McMillan, 1999       0.6               linear
E. P. wisneri (Kuo, Hang and       0.5-0.7           fairly straight
  Mok, 1994)
E. Q. yangi# (Teng, 1958)          0.8-1.0           crowded

Species                            Last GA and PCD     Eyespots

E. Q. ancon Mok. Saavedra-Diaz &   separate            present
  Acero-P, 2001
E. P. atami (Dean, 1904)           fused               faint or absent
E. E. burgeri# (Girard, 1855)      fused (90%)         present
E. P. cheni# (Shen and Tao, 1975)  separate            absent
E. E. bischoffli (Schneider,       separate            present
  1880)
E. E. caribbeaus Fernholm, 1982    --                  --
E. E. carlhbbsi McMillan and       fused               present
  Wisner, 1984
E. E. chinensis# Kuo and Mok,      fused               present
  1994
E. E. cirrhatus# (Forster, 1801)   fused (exceptions)  present
E. E. deani# (Evermann and         fused               present
  Goldsborough, 1907)
E. E. eos Fernholm, 1991           fused               absent
E. E. fernholmi McMillan and       fused               absent
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and     separate            absent
  Mok, 1994)
E. E. fritzi Wisner and            fused               present
  McMillan, 1990
E. E. goliath Mincarone and        fused               present
  Stewart, 2006
E. E. grouseri McMillan, 1999      fused               prominent
E. E. hexatrema (Muller, 1836)     fused               --
E. E. indrambaryai Wongratana,     fused               present
  1983
E. E. lakeside Mincarone and       fused               weak
  McCosker, 2004
E. E. laurahubbsae McMillan and    fused               weak
  Wisner, 1984
E. E. longipinnis# Strahan, 1975   fused               absent
E. E. mcconnaugheyi Wisner and     fused               present
  McMillan, 1990
E. E. mccoskeri McMillan, 1999     fused               weak
E. E. mendozai Hensley, 1985       fused               present
E. E. menezesi Mincarone, 2000     --                  present
E. E. minor Fernholm and Hubbs,    fused               --
  1981
E. P. moki (McMillan and Wisner,   fused               faint or absent
  2004)
E. E. mulitidens Fernholm and      fused               present
  Hubbs, 1981
E. E. nanii Wisner and McMillan,   separate (61%)      present
  1988
E. Q. nelsoni# (Kuo, Huang and     fused               absent
  Mok, 1994)
E. E. octatrema (Barnard, 1923)    fused               --
E. E. okinoseanus (Dean, 1904)     fused               present
E. E. polytrema (Girard, 1855)     fused (91%)         present
E. E. profundus (Barnard, 1923)    --                  --
E. P. sheni# (Kuo, Huang and       fused               prominent
  Mok, 1994)
E. E. sinus Wisner and McMillan,   fused               present
  1990
E. P. springeri (Bigelow and       --                  --
  Schroeder, 1952)
E. E. stoutii# (Lockington, 1878)  fused               present
E. E. strahani McMillan and        fused               absent
  Wisner, 1984
E. E. strickrotti# n. sp.          separate            absent
E. Q. taiwanae# (Shen and Tao,     fused               faint or absent
  1975)
E. P. walkeri (McMillan and        fused               faint or absent
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz     fused               present
  and Acero, 2001
E. E. wisneri McMillan, 1999       fused               distinct
E. P. wisneri (Kuo, Hang and       fused               present
  Mok, 1994)
E. Q. yangi# (Teng, 1958)          fused               absent

Species                            Color                      Max TL

E. Q. ancon Mok. Saavedra-Diaz &   light brown                 220
  Acero-P, 2001
E. P. atami (Dean, 1904)           purplish-brown-grey         610
E. E. burgeri# (Girard, 1855)      brownish                    690
E. P. cheni# (Shen and Tao, 1975)  greyish brown               386
E. E. bischoffli (Schneider,       black-brown                 550
  1880)
E. E. caribbeaus Fernholm, 1982    light tan                   385
E. E. carlhbbsi McMillan and       dark brown                 1160
  Wisner, 1984
E. E. chinensis# Kuo and Mok,      grey brownish               540
  1994
E. E. cirrhatus# (Forster, 1801)   dark brown, white marks     830
E. E. deani# (Evermann and         black (rarely pinkish)      523
  Goldsborough, 1907)
E. E. eos Fernholm, 1991           pink                        665
E. E. fernholmi McMillan and       brownish                    373
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and     greyish brown               359
  Mok, 1994)
E. E. fritzi Wisner and            purplish black              592
  McMillan, 1990
E. E. goliath Mincarone and        dark brown                 1275
  Stewart, 2006
E. E. grouseri McMillan, 1999      ivory to brownish-black     420
E. E. hexatrema (Muller, 1836)     light-blackish brown        720
E. E. indrambaryai Wongratana,     purplish-brown              437
  1983
E. E. lakeside Mincarone and       pink-orange                 275
  McCosker, 2004
E. E. laurahubbsae McMillan and    --                          375
  Wisner, 1984
E. E. longipinnis# Strahan, 1975   brownish                    550
E. E. mcconnaugheyi Wisner and     brownish                    470
  McMillan, 1990
E. E. mccoskeri McMillan, 1999     brownish-black              320
E. E. mendozai Hensley, 1985       bluish-grey                 450
E. E. menezesi Mincarone, 2000     light brown                 737
E. E. minor Fernholm and Hubbs,    --                          395
  1981
E. P. moki (McMillan and Wisner,   dark brown                  470
  2004)
E. E. mulitidens Fernholm and      dark                        815
  Hubbs, 1981
E. E. nanii Wisner and McMillan,   black-brown                 664
  1988
E. Q. nelsoni# (Kuo, Huang and     brownish-grey               234
  Mok, 1994)
E. E. octatrema (Barnard, 1923)    brownish                    300
E. E. okinoseanus (Dean, 1904)     brownish                    800
E. E. polytrema (Girard, 1855)     piebald-dark                460
E. E. profundus (Barnard, 1923)    dark brown                  620
E. P. sheni# (Kuo, Huang and       dark brown                  436
  Mok, 1994)
E. E. sinus Wisner and McMillan,   reddish brown               481
  1990
E. P. springeri (Bigelow and       greyish brown               590
  Schroeder, 1952)
E. E. stoutii# (Lockington, 1878)  brownish (rarely pinkish)   468
E. E. strahani McMillan and        brownish                    520
  Wisner, 1984
E. E. strickrotti# n. sp.          pink                        314
E. Q. taiwanae# (Shen and Tao,     brownish-grey               334
  1975)
E. P. walkeri (McMillan and        brownish                    518
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz     pink                        216
  and Acero, 2001
E. E. wisneri McMillan, 1999       brownish-black              356
E. P. wisneri (Kuo, Hang and       light brown                 388
  Mok, 1994)
E. Q. yangi# (Teng, 1958)          brownish-grey               296

Species in bold included in the molecular phylogeny.
Abreviations: E. = Eptatretus; Q. = Quadratus; P. = Paramyxine; NSP =
nasal sinus papillae; GP = gill pouches; DM = dental muscle; ABA =
afferent branchial arteries; MVA = median ventral aorta; VA = ventral
aorta; PCD = pharyngocutaneous duct; GA = gill apertures; VFF = ventral
fin fold. Blanks as (--) where an extensive literature search faild to
provide data.

Note: Species indicated with # included in the molecular phylogeny.

Appendix Table 2. Body measurements of Eptatretus spp. in % of total
length (TL)

                                 Prebr.     Branchial  Trunk   Tail
Species                          length     length     length  length

E. Q. ancon Mok. Saavedra-       37          1.95      48      12.0
  Diaz & Acero-P, 2001
E. P. atami (Dean, 1904)         27-30       1.3-4.2   54-56   11.1-14.2
E. E. burgeri# (Girard, 1855)    25-30       6.2-7.8   48-55   13-17
E. P. cheni# (Shen and Tao,      33-36       2.2-3.4   46-51   13.2-16.7
  1975)
E. E. bischoffli (Schneider,     17.6-22.4  11.4-16.1  45-54   14.0-21.8
  1880)
E. E. caribbeaus Fernholm, 1982  21-24       5.8-7.8   50-56   16.5-19.6
E. E. carlhubbsi McMillan and    17-20       5.5-7.7   58-62   14.5-17.6
  Wisner, 1984
E. E. chinensis# Kuo and Mok,    25-35       4.4-6.4   46-53   13.9-18.6
  1994
E. E. cirrhatus# (Forster,       21-24       6.9-8.9   53-56   13.5-16.8
  1801)
E. E. deani# (Evermann and       14-20      12.7-18.2  48-56   12.6-19.2
  Goldsborough, 1907)
E. E. eos Fernholm, 1991         23.5        4.7       54      18.0
E. E. fernholmi McMillan and     21          8.0       56      14.9
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and   30-33       2.0-2.9   50-52   13.4-16.3
  Mok, 1994)
E. E. fritzi Wisner and          18-25      11.5-15.8  46-56   13.2-18.1
  McMillan, 1990
E. E. goliath Mincarone and      18.8        6.7       58.8    15.7
  Stewart, 2006
E. E. grouseri McMillan, 1999    20-24       6.3-8.1   54-57   14.6-17.5
E. E. hexatrema (Muller, 1836)   27-32       5-6       --      --
E. E. indrambaryai Wongratana,   21-23       8.6-10.6  52-57   15.8-18.4
  1983
E. E. lakeside Mincarone and     25          6.2       51      18.2
  McCosker, 2004
E. E. laurahubbsae McMillan and  18-20       5.2-5.9   55-59   18.1-21.3
  Wisner, 1984
E. E. longipinnis Strahan,       29          4.0       59       8.3
  1975#
E. E. mcconnaugheyi Wisner and   15-18      16-22      48-56   12.2-16.1
  McMillan, 1990
E. E. mccoskeri McMillan, 1999   24-26       9.3-10.1  49-50   15.6-17.7
E. E. mendozai Hensley, 1985     22-25       4.7-6.6   51-55   16.2-19.3
E. E. menezesi Mincarone, 2000   19-27       5.0-8.3   51-65   14.6-22.0
E. E. minor Fernholm and Hubbs,  20-26       5.1-7.2   51-56   13.9-18.3
  1981
E. P. moki (McMillan and         27-31       1.4-3.0   49-55   15.4-19.5
  Wisner, 2004)
E. E. mulitidens Fernholm and    19-21       6.1-6.9   55-57   16.9-18.8
  Hubbs, 1981
E. E. nanii Wisner and           12.8-15.6  17.5-22.0  48-53   15.2-17.3
  McMillan, 1988
E. Q. nelsoni# (Kuo, Huang and   31-33       1.1-2.8   50-53   15.0-18.0
  Mok, 1994)
E. E. octatrema (Barnard, 1923)  25         --         --      --
E. E. okinoseanus (Dean, 1904)   19-23       6.2-9.2   50-59   12.7-15.5
E. E. polytrema (Girard, 1855)   13.9-16.9  16.8-20.1  48-53   12.7-17.9
E. E. profundus (Barnard, 1923)  20          6.0       55      --
E. P. sheni# (Kuo, Huang and     25-31       2.4-4.2   53-56   14.4-16.7
  Mok, 1994)
E. E. sinus Wisner and           20-28       9.2-17.2  45-54   10.2-17.4
  McMillan, 1990
E. P. springeri (Bigelow and     22-27       2.5-5.6   53-61   13.4-16.8
  Schroeder, 1952)
E. E. stoutii# (Lockington,      19-25      11.5-14.2  47-54   10.4-17.8
   1878)
E. E. strahani McMillan and      21-23       5.2-8.7   48-56   17.4-20.2
  Wisner, 1984
E. E. strickrotti# n. sp.        19.7        9.3       58.9    12.1
E. Q. taiwanae# (Shen and Tao,   28-35       1.3-2.7   52-56   12.1-14.6
  1975)
E. P. walkeri (McMillan and      27-32       2.0-3.1   53-58   13.0-15.6
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz   31-33       3.0-3.5   48-50   10.9-14.9
  and Acero, 2001
E. E. wisneri McMillan, 1999     19-23      10.0-11.1  50-53   16.7-17.1
E. P. wisneri (Kuo, Hang and     28-31       4.6       49.4    13.6
  Mok, 1994)
E. Q. yangi# (Teng, 1958)        29-32       1.1-1.7   53-55   12.2-15.6

                                 Depth      Depth at
Species                          w/o VFF    cloaca    Tail depth

E. Q. ancon Mok. Saavedra-        6.8       --         7.0
  Diaz & Acero-P, 2001
E. P. atami (Dean, 1904)          7.9-8.0   6.3-6.8    7.4-8.8
E. E. burgeri# (Girard, 1855)     4.0-8.3   4.5-5.8    5.1-8.5
E. P. cheni# (Shen and Tao,       7.7-9.9   7.8-9.9    7.6-10.2
  1975)
E. E. bischoffli (Schneider,      7.1-11.2  5.6-8.7    5.6-8.3
  1880)
E. E. caribbeaus Fernholm, 1982   7.4-10.6  9.6-9.2    7.5-10.9
E. E. carlhubbsi McMillan and     7.8-10.6  6.5-8.5    8.9-10.5
  Wisner, 1984
E. E. chinensis# Kuo and Mok,    --         --         7.1-10.2
  1994
E. E. cirrhatus# (Forster,        8.1-10.2  5.7-7.4    7.7-9.1
  1801)
E. E. deani# (Evermann and        4.5-10.5  3.8-8.5    5.2-10.3
  Goldsborough, 1907)
E. E. eos Fernholm, 1991         --         --        --
E. E. fernholmi McMillan and      8.0       6.0        8.6
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and    5.6-8.0   5.2-6.3    7.0-10.5
  Mok, 1994)
E. E. fritzi Wisner and           5.3-10.2  4.5-7.2    5.8-9.2
  McMillan, 1990
E. E. goliath Mincarone and       9.5       8.2       11.4
  Stewart, 2006
E. E. grouseri McMillan, 1999     4.2-8.8   5.7-6.5    6.3-7.9
E. E. hexatrema (Muller, 1836)    6-7       --        --
E. E. indrambaryai Wongratana,    8.5-10.3  6.0-8.9    7.6-9.9
  1983
E. E. lakeside Mincarone and      6.4       5.4        6.0
  McCosker, 2004
E. E. laurahubbsae McMillan and   7.3-9.1   6.1-8.0    8.2-8.9
  Wisner, 1984
E. E. longipinnis Strahan,        4.0       3.5        4.2
  1975#
E. E. mcconnaugheyi Wisner and    5.5-9.0   4.7-7.1    6.0-8.8
  McMillan, 1990
E. E. mccoskeri McMillan, 1999    9.4-10.6  7.3        8.7-10.2
E. E. mendozai Hensley, 1985      8.0-10.9  6.5-8.2    8.0-9.4
E. E. menezesi Mincarone, 2000    7.1-10.6  6.6-9.1    6.6-10.6
E. E. minor Fernholm and Hubbs,   7.1-10.8  5.2-7.9    5.3-11.6
  1981
E. P. moki (McMillan and          5.2-8.2   5.8-8.2    7.7-9.7
  Wisner, 2004)
E. E. mulitidens Fernholm and     7.8-11.3  6.0-8.1    6.6-8.6
  Hubbs, 1981
E. E. nanii Wisner and            7.3-9.9   4.2-7.6    6.2-9.2
  McMillan, 1988
E. Q. nelsoni# (Kuo, Huang and   14.7-15.3  --         8.9-10.1
  Mok, 1994)
E. E. octatrema (Barnard, 1923)   4.0       --        --
E. E. okinoseanus (Dean, 1904)    5.5-7.9   6.0        6.2-9.0
E. E. polytrema (Girard, 1855)    6.8-10.9  4.9-8.0    5.7-8.6
E. E. profundus (Barnard, 1923)   8.3       --        --
E. P. sheni# (Kuo, Huang and      6.9-9.2   --         8.3-10.0
  Mok, 1994)
E. E. sinus Wisner and            4.6-10.1  3.9-8.6    4.8-9.0
  McMillan, 1990
E. P. springeri (Bigelow and      6.2-9.7   --         6.4-9.3
  Schroeder, 1952)
E. E. stoutii# (Lockington,       4.1-9.0   3.8-7.9    4.5-8.3
   1878)
E. E. strahani McMillan and      10.1-12.5  --        10.9-12.5
  Wisner, 1984
E. E. strickrotti# n. sp.         2.9       2.6        2.4
E. Q. taiwanae# (Shen and Tao,    6.3-10.6  --         8.1-11.8
  1975)
E. P. walkeri (McMillan and       6.2-8.2   5.6-6.8    7.1-8.3
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz    7.0-7.7   --        11.0-10.1
  and Acero, 2001
E. E. wisneri McMillan, 1999      8.5-8.9   6.5        7.6-7.8
E. P. wisneri (Kuo, Hang and      9.3       8.1       10.5
  Mok, 1994)
E. Q. yangi# (Teng, 1958)         6.3-9.6   6.3-8.8    6.5-10.0

Species                          Locality                    Depth m

E. Q. ancon Mok. Saavedra-       Caribbean Sea                470-488
  Diaz & Acero-P, 2001
E. P. atami (Dean, 1904)         off Japan                    300-536
E. E. burgeri# (Girard, 1855)    off Japan, Korea, China &     10-270
                                   Taiwan
E. P. cheni# (Shen and Tao,      off SW Taiwan                180-268
  1975)
E. E. bischoffli (Schneider,     off Chile                      8-50
  1880)
E. E. caribbeaus Fernholm, 1982  Caribbean Sea                365-500
E. E. carlhubbsi McMillan and    off Hawaii, Wake             481-1574
  Wisner, 1984                     Isl., Guam
E. E. chinensis# Kuo and Mok,    South China Sea              500-600
  1994
E. E. cirrhatus# (Forster,       off New Zealand, S & E        30-700
  1801)                            Australia
E. E. deani# (Evermann and       NE Pacific, off N.           107-2743
  Goldsborough, 1907)              America
E. E. eos Fernholm, 1991         Tasman Sea                   991-1013
E. E. fernholmi McMillan and     off Philippines              560
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and   off Taiwan                   300-412
  Mok, 1994)
E. E. fritzi Wisner and          off Guadeloupe Isl.,          18-2743
  McMillan, 1990                   Mexico
E. E. goliath Mincarone and      off N New Zealand            811
  Stewart, 2006
E. E. grouseri McMillan, 1999    off Galapagos                722
E. E. hexatrema (Muller, 1836)   off South Africa              10-400
E. E. indrambaryai Wongratana,   Andaman Sea                  267-400
  1983
E. E. lakeside Mincarone and     off Galapagos                762
  McCosker, 2004
E. E. laurahubbsae McMillan and  SE Pacific, off Chile       2400
  Wisner, 1984
E. E. longipinnis Strahan,       off South Australia           40
  1975#
E. E. mcconnaugheyi Wisner and   Gulf of California            42-415
  McMillan, 1990
E. E. mccoskeri McMillan, 1999   off Galapagos                215
E. E. mendozai Hensley, 1985     Caribbean Sea                720-1100
E. E. menezesi Mincarone, 2000   SW Atlantic, off Brazil      250-530
E. E. minor Fernholm and Hubbs,  Gulf of Mexico               300-400
  1981
E. P. moki (McMillan and         off Japan                    100
  Wisner, 2004)
E. E. mulitidens Fernholm and    Caribbean Sea and NE         239-770
  Hubbs, 1981                      Brazil
E. E. nanii Wisner and           off Chile                    274
  McMillan, 1988
E. Q. nelsoni# (Kuo, Huang and   off SW Taiwan                 50-858
  Mok, 1994)
E. E. octatrema (Barnard, 1923)  off South Africa              45-75
E. E. okinoseanus (Dean, 1904)   off Japan, Taiwan            300-1020
E. E. polytrema (Girard, 1855)   off Chile                     10-350
E. E. profundus (Barnard, 1923)  off South Africa             732
E. P. sheni# (Kuo, Huang and     off SW Taiwan                200-619
  Mok, 1994)
E. E. sinus Wisner and           Gulf of California           198-1330
  McMillan, 1990
E. P. springeri (Bigelow and     Gulf of Mexico               400-730
  Schroeder, 1952)
E. E. stoutii# (Lockington,      NE Pacific, off N. America    16-633
   1878)
E. E. strahani McMillan and      South China Sea,             189
  Wisner, 1984                     Philippines
E. E. strickrotti# n. sp.        East Pacific Rise           2211
E. Q. taiwanae# (Shen and Tao,   off NE Taiwan                 20-427
  1975)
E. P. walkeri (McMillan and      off Japan                     75-120
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz   Caribbean Sea                300-306
  and Acero, 2001
E. E. wisneri McMillan, 1999     off Galapagos                563
E. P. wisneri (Kuo, Hang and     off SE Taiwan                200
  Mok, 1994)
E. Q. yangi# (Teng, 1958)        off NE and SW Taiwan          20-225

Species                          Major sources of information

E. Q. ancon Mok. Saavedra-       Mok et al., 2001
  Diaz & Acero-P, 2001
E. P. atami (Dean, 1904)         McMillan and Wisner, 2004
E. E. burgeri# (Girard, 1855)    McMillan and Wisner, 2004
E. P. cheni# (Shen and Tao,      McMillan and Wisner, 2004
  1975)
E. E. bischoffli (Schneider,     Wisner and McMillan, 1988
  1880)
E. E. caribbeaus Fernholm, 1982  Fernholm, 1982; Mincarone, 2000
E. E. carlhubbsi McMillan and    McMillan and Wisner, 1984
  Wisner, 1984
E. E. chinensis# Kuo and Mok,    Chen et al., 2005; McMillan and
  1994                             Wisner, 2004
E. E. cirrhatus# (Forster,       McMillan and Wisner, 1984, present
  1801)                            study
E. E. deani# (Evermann and       Wisner and McMillan, 1990
  Goldsborough, 1907)
E. E. eos Fernholm, 1991         Fernholm, 1991; Mok et al., 2001
E. E. fernholmi McMillan and     McMillan and Wisner, 2004
  Wisner, 2004
E. P. fernholmi (Kuo, Hang and   McMillan and Wisner, 2004
  Mok, 1994)
E. E. fritzi Wisner and          Wisner and McMillan, 1990
  McMillan, 1990
E. E. goliath Mincarone and      Mincarone and Stewart, 2006
  Stewart, 2006
E. E. grouseri McMillan, 1999    McMillan, 1999; Mincarone and
                                   McCosker, 2004
E. E. hexatrema (Muller, 1836)   Fernholm, 1986; Strahan, 1975
E. E. indrambaryai Wongratana,   Wongratana, 1983
  1983
E. E. lakeside Mincarone and     Mincarone and McCosker, 2004
  McCosker, 2004
E. E. laurahubbsae McMillan and  McMillan and Wisner, 1984
  Wisner, 1984
E. E. longipinnis Strahan,       Strahan, 1975; present study
  1975#
E. E. mcconnaugheyi Wisner and   Wisner and McMillan, 1990
  McMillan, 1990
E. E. mccoskeri McMillan, 1999   McMillan, 1999
E. E. mendozai Hensley, 1985     Hensley, 1985; Mok et al, 2001
E. E. menezesi Mincarone, 2000   Mincarone, 2000
E. E. minor Fernholm and Hubbs,  Fernholm and Hubbs, 1981
  1981
E. P. moki (McMillan and         McMillan and Wisner, 2004
  Wisner, 2004)
E. E. mulitidens Fernholm and    Fernholm and Hubbs, 1981; Mincarone
  Hubbs, 1981                      and Sampaio, 2004
E. E. nanii Wisner and           Wisner and McMillan, 1988
  McMillan, 1988
E. Q. nelsoni# (Kuo, Huang and   McMillan and Wisner, 2004
  Mok, 1994)
E. E. octatrema (Barnard, 1923)  Barnard, 1923; Fernholm, 1986
E. E. okinoseanus (Dean, 1904)   McMillan and Wisner, 2004
E. E. polytrema (Girard, 1855)   Wisner and McMillan, 1988; Wisner,
                                   1999
E. E. profundus (Barnard, 1923)  Barnard, 1923; Strahan, 1975;
                                   Mincarone and McCosker, 2004
E. P. sheni# (Kuo, Huang and     McMillan and Wisner, 2004
  Mok, 1994)
E. E. sinus Wisner and           Wisner and McMillan, 1990
  McMillan, 1990
E. P. springeri (Bigelow and     Mok et al., 2001
  Schroeder, 1952)
E. E. stoutii# (Lockington,      Wisner and McMillan, 1990
   1878)
E. E. strahani McMillan and      McMillan and Wisner, 2004
  Wisner, 1984
E. E. strickrotti# n. sp.        Present study
E. Q. taiwanae# (Shen and Tao,   McMillan and Wisner, 2004; Mok et al.,
  1975)                            2001
E. P. walkeri (McMillan and      McMillan and Wisner, 2004
  Wisner, 2004)
E. E. wayun Mok, Saavedra-Diaz   Mok et al., 2001
  and Acero, 2001
E. E. wisneri McMillan, 1999     McMillan, 1999
E. P. wisneri (Kuo, Hang and     McMillan and Wisner, 2004
  Mok, 1994)
E. Q. yangi# (Teng, 1958)        McMillan and Wisner, 2004; Mok et al.,
                                   2001

Species in bold included in the molecular phylogeny.
Abbreviations: E = Eptatretus; Q = Quadratus; P = Paramyxine.

Note: Species indicated with # included in the molecular phylogeny.


PETER R. MOLLER (1,*) AND W. JOE JONES (2)

(1) Zoological Museum, Natural History Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen East, Denmark; and (2) Monterey Bay Aquarium Research Institute (MBARI), Moss Landing, California 95039-9644

Received 21 April 2006; accepted 21 November 2006.

* To whom correspondence should be addressed. E-mail: pdrmoller@snm.ku.dk

Abbreviations: BPP, Bayesian posterior probabilities; E, Eptatretus; GA, gill aperture; GP, gill pouche; P, Paramyxine; PCD, pharyngocutaneous duct; Q, Quadratus; TL, total length; VA, ventral aorta.
Table 1 Summary of information for the specimens used for sequencing of
16S rRNA or phylogenetic analyses

Species                    Voucher No.  Locality           GenBank No.

Eptatretus E. burgeri      --           Taiwan             AF364617,
                                                             AY619579,
                                                             AF364616
Eptatretus P. cheni        --           Taiwan             AF364620-1
Eptatretus E. chinensis    --           Taiwan             AY619580
Eptatretus E. cirrhatus    --           New Zealand        AF364619
Eptatretus E. deani        KU 28249     California, USA    EF014477
Eptatretus E. longipinnis  SAMA F07540  South Australia    EF014476
Eptatretus Q. nelsoni      --           Taiwan             AF364608
Eptatretus P. sheni        --           Taiwan             AF364610
Eptatretus E. stoutii      --           Oregon, USA        AF364618
Eptatretus E. strickrotti  CAS 223480   East Pacific Rise  EF014478
  n. sp.
Eptatretus E. taiwanae     --           Taiwan             AF364611
Eptatretus Q. yangi        --           Taiwan             AF364612-5
Myxine circifrons          --           Unknown            AF364628-9
Myxine formosana           --           Taiwan             AF364625
Myxine sp. 1               --           Taiwan             AF364622-4
Myxine sp. 2               --           Taiwan             AF364626
Myxine sp. 3               --           Taiwan             AF364627
Petromyzon marinus         --           Unknown            U11880

Dashes (--) indicate that data were not available from Kuo et al.
(2003).
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Author:Moller, Peter R.; Jones, W. Joe
Publication:The Biological Bulletin
Article Type:Author abstract
Geographic Code:0PACR
Date:Feb 1, 2007
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