Eptatretus strickrotti n. sp. (Myxinidae): first hagfish captured from a hydrothermal vent.
IntroductionThe 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 |
| Words: | 9511 |
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