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Deep-sea Benthic Fish of the Hawaiian Archipelago, Cross Seamount, and Johnston Atoll! PDF

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PacificScience(1994), vol. 48, no. 4: 367-409 © 1994byUniversityofHawaiiPress. All rights reserved Deep-sea Benthic Fish ofthe Hawaiian Archipelago, Cross Seamount, and Johnston Atoll! E. H. CHAVE2AND B. C. MUNDy3 ABSTRACT: More than 250 benthic fish taxa were photographed and video taped by Hawaii Undersea Research Laboratory submersibles at depths be tween 40 and 2000 m in the Hawaiian Archipelago, Johnston Atoll, and Cross Seamount. Most ofthe 213 identified fish species occurred close to hard sub strates with holes, ledges, or caves. Twenty-two species (notably the larger sharks, lutjanids, and carangids) are cosmopolitan. Seventy-six species are restricted to various Indo-Pacific areas, 64 in the Pacific, and 51 in the Hawaiian Archipelago including Cross Seamount and Johnston Atoll. There is a rapid decrease in the number of species from 200 to 400 m depth. One hundred eight species were seen 20 m deeper than previously reported. Eleven of the deeper-dwelling animals were found 20 m shallower than pre viouslyrecorded. Faunalzoneswerenotrecognizedatanydepth. Speciesnewly recorded in Hawai'i include Bathypterois grallator (Goode & Bean), Bodianus cylindriatus (Tanaka), Centrophorus cf. granulosus (Bloch & Schneider), Chau naxfimbriatus Hilgendorf, Caelorinchus spilonotus Sazonov & Iwamoto, Noto canthus sp., Paratrachichthysprosthemius Jordan & Fowler, Prognathodes gue zei (Mauge & Bauchot), and Sladenia remiger Smith & Radcliffe. New species collectedandreportedelsewhereare CentrodracorubellusFrickeetaI.,Epigonus glossodontusGon, Owstoniasp., andPseudanthiasfucinus(Randall & Ralston). Caelorinchus sp. 2and Callanthiassp. areprobablyundescribed. Itappears that the Hawaiian deep-sea fish fauna has multiple origins and affinities with many regions. THIS PAPER SUMMARIZES the fish fauna col waiian Archipelago (160 42' N, 1690 32' W). lected, photographed, and videotaped during The data were obtained from dives made at 500 dives conducted by Hawaii Undersea depths between 40 and 2000 m from 1982 to Research Laboratory (HURL) submersibles 1992. Makali'i and Pisces V. These vehicles oper The deep-sea fish fauna ofHawai'i is now ated in the Hawaiian Archipelago from among the best known in the Pacific, when French Frigate Shoals to L6'ihi Volcano, the results of the trawl surveys by Gilbert located off the southeastern end of Hawai'i (1905) and Struhsaker (1973) are combined Island (180 55' N, 1550 16' W); on Cross with results ofthis submersible study. Trawl Seamount, situated southwest of Hawai'i studies are limited by factors such as net Island (180 44' N, 1580 15' W); and at John avoidance and imprecise knowledge ofdepth ston Atoll, located southwest of the Ha- of capture (Haedrich et ai. 1975, Merrett and Marshall 1981). Submersible studies are biased by attraction or repulsion of organ 1Manuscriptaccepted II December1993. isms and photographic selectivity of the 2Hawaii Undersea Research Laboratory, University scientists (Ralston et ai. 1986, Chave and ofHawaii,Honolulu, Hawai'i96822. Jones 1991; HURL records, unpubi. data). 3Honolulu Laboratory, Southwest Fisheries Science Many of the sampling limitations are com Center, National Marine Fisheries Service, NOAA, Honolulu, Hawai'i96822. plementary when precise submersible ob- 367 FIGURE1. Hexanchus griseus (HURL photo 43-31, Colin). FIGURE4. Echinorhinus cookei (HURL photo 5039 43, Mullineaux). FIGURE5. Torpedo sp. (HURL photo 119-68, Chave). Figure2. Pseudotriakis microdon (HURL video 5050,Polovina). Figure3. Squalus mitsukurii (HURL photo 148-94, Figure6. Hexatrygon longirostra (HURL photo Chave). 5069-06,Moore). Deep-seaFish ofthe Hawaiian Archipelago-CHAVE ANDMUNDY 369 FIGURE8. Hydrolagus purpurescens (HURL photo 238-11, France). FIGURE7. Plesiobatis daviesi (HURL photo 87-18, Maragos). FIGURE9. Anilyophineeelandthezoanthid Gerardiasp. (HURLphoto68-08,Chave). 370 PACIFICSCIENCE,Volume48, October 1994 Figure12. Gadomus melanopterus (HURL photo FIGURE10. Nettastoma parviceps (HURL photo 5091-76,Malahoff). 5068-11,Moore). FIGURE11. Chlorophthalmus proridens (HURL Figure13. Caelorinchus sp. 2 (HURL video 5204, photo123-13,Chave). Young). servations of depth and behavior are com MATERIALS AND METHODS bined with data on large numbers of speci mens and areas sampled by trawls. We can The submersibles' still and video cameras address the biogeography, depth distri were synchronized to record the same butions, and ecology of Hawaiian deep-sea images, enabling subjects seen on the video fish with greater confidence than previously screen to be photographed. Photos were possible. taken in strobe light. The videocamera ran FIGURE17. Gadella molokaiensis (HURL photo Figure14. Coryphaenoides longicirrhus (HURL 118-25,Chave). video5180, Malahoff). FIGURE15. Nezumia propinqua (HURL photo 215 46, Chave). Figure18. Pycnocraspedum armatum (HURL photo 5068-49, Moore). FIGURE19. Lophiodes miacanthus (HURL photo 144-17,Scheuer). FIGURE16. Antimora microlepis (HURL photo 5145-88,Garcia). 372 PACIFIC SCIENCE, Volume48, October 1994 FIGURE22. Ijimaiaplicatellus(HURLphoto 34-146, Colin). FIGURE20. Sladenia remiger (HURL photo 5186 141,Garcia). FIGURE23. Beryx decadactylus (HURL photo 215 05,Chave). istry profiles, depth records, and speCImens FIGURE21. Malthopsis jordani (HURL video 2025, augment these data. Young). All specimens and many of the photo graphs were sentto experts for identification. Some identifications are tentative because throughout the dive, much ofthe time using specimens were not collected or because the the submersible's running lights for illumina photographs did not show those character tion. The floodlights were turned to video istics used for identification. The photo tape selected subjects. All photographs from graphs and videotapes documenting the fish depths greater than 40 m were examined. species in this study are filed in the HURL About 2100 ofthese include identifiable fish archives and are available on request. Fig and form the data set used in this study. ures 1-45 and Plates I-IV contain the first Videotapes, lightmeasurements, waterchem- published underwater photographs of some Deep-seaFishofthe Hawaiian Archipelago-CHAVE AND MUNDY 373 FIGURE26. Zenopsis nebulosus (HURL photo 357 FIGURE24. Polymixiaberndti(HURLphoto213-48, 37,Grigg). Chave). FIGURE25. Stethopristis eos (HURL photo 155-13, Chave). of the fishes. Previously published photos taken from HURL submersibles may be FIGURE27. Grammicolepis brachiusculus (HURL video 119,Chave). found in Randall and Ralston (1984), Ran dall etal. (1985), Ralston et al. (1986), Jones and Sulak (1990), Chave and Jones (1991), relative abundances. Table 2 contains new and Pyle and Chave (1994). information about some of the fish. The Table 1 includes all identified fish taxa, order of species presentation in both tables their depth ranges, locations observed, and follows Eschmeyer (1990). 374 PACIFICSCIENCE,Volume 48, October 1994 FIGURE28. Bembradium roseum (HURL photo 130 28,Maragos). FIGURE30. Symphysanodonmauna/oae(HURLvideo 272,Grigg). Bean], Bodianus cylindriatus [Tanaka], Cae lorinchus spilonotus Sazonov & Iwamoto, Centrophorus cf. granulosus [Bloch & Schneider], Chaunax fimbriatus Hilgendorf, Notocanthus sp., Paratrachichthys prosthe mius Jordan & Fowler, Prognathodes guezei [Mauge & Bauchot], and Sladenia remiger Smith & Radcliffe) are also discussed in Table 2. BehaviorandMorphology Below 100m mostfish were located on or Figure29. Synagropsargyrea(HURLphoto 118-18, just above the bottom. This agrees with ob Chave). servations of deep-sea benthic fish seen pri marily at 0-5 m from the bottom on the Nazca and Sala-y-G6mez Ridges (Golovan and Pakhorukov 1987). Most species were RESULTS AND DISCUSSION found on or near hard substrates that Mostfish seen in this study were collected containedholes, caves, orotherirregularities. previously in the Pacific (Table 1). Collec On sand, large groups of fish were seen in tions made with the HURL submersibles troughs when the currents were strong. They yielded four new species: Centrodraco ru were only observed above the substratewhen bel/us Fricke et al., Epigonus glossodontus currents swept particles through the water at Gon, Owstonia sp., and Pseudanthiasfucinus moderate rates (HURL records, unpubl. (Randall & Ralston) (Table 2). Two possible data). new species (Caelorinchus sp. 2 and Cal/an When the submersibles were operating thias sp.) and nine other new Hawaiian withtheirrunninglightson, mostfish did not records (Bathypterois gral/ator [Goode & react to the vehicles. Fishes thatdid reactare Deep-seaFish ofthe HawaiianArchipelago-CRAVE AND MUNDY 375 FIGURE31. AnaggregationofSymphysanodon typus(HURLphoto337-48,Ralston). moving toward or away from the light, or becomingmotionless. Benthicfish deeper than 100m were com monly eel-like in shape. Many could not be identified from their photos. Ate1eopids, macrourids, halosaurids, and congrids were often the most abundant of the eel-like forms. Perhaps this body shape offers less re sistance to current flow and allows greater motility for near-bottom animals (Zaferman 1992). Having a long lateral line also may increase sensory capabilities and enable ee1 likefish to search for preywithoutdisturbing the bottom (Marshall 1971, Marshall and Merrett 1977). FIGURE32. Eumegistus illustris (HURL photo 5070 16,Moore). Biogeography Of 213 species of fish identified in this listed in Table 2. When the brighter photo study and listed in Table 1, 25 have circum and video lights were turned on, many ofthe global distributions, 73 are found in the fish that occurred deeper than 100m reacted Indo-Pacific, and 64 are restricted to Pacific by bumping into the bottom, backing away, waters. Fifty-one species (24%) are found 376 PACIFICSCIENCE,Volume 48, October 1994 FIGURE33. Aphareusrutilans(bottomright) andseveral Pristipomoidesfilamentosus(HURLphoto 336-03, Ral ston). waiian Islands (HURL records [unpubl.]; Grigg et al. 1987). This observation was ex pected because Cross Seamount lies only 190 km to the west and L6'ihi is 50 km to the south of Hawai'i Island, easily within the range oflarval fish dispersal. Two trends are evident from distribution datainTable 1. Mostlarge, strongswimmers with carnivorous or omnivorous habits (e.g., many ofthe sharks, lutjanids, and carangids) have ocean-wide distributions. Fish found shallowerthan 100m tend to have more spe cific habitat preferences and wider zoogeo graphic distributions than deeper-dwelling fish, perhaps because shallower species have Figure34. Etelis carbunculus(HURL photo 213-74, been studiedmore extensively. Chave). Two hypotheses are currently recognized as explaining the distribution of shallow watermarine organismsin Hawai'i: dispersal only in the Hawaiian Archipelago, Cross and vicariance. Both are discussed at length Seamount, and Johnston Atoll. This high by Springer (1982) and Newman (1986). number of endemic deep-sea species agrees The dispersal hypothesis assumes that the with calculations that include most of the Hawaiian Archipelago was colonized by shallow-water fish in Central Pacific areas larval drift of Indo-Pacific fish across "filter (Randall 1985, Randall et al. 1985, Kosaki bridges" (Gosline 1955, Gosline and Brock et al. 1991). All fish seen on Cross Seamount 1960) or along "stepping stones" (Wilson and L6'ihi were also observed off the Ha- and Kaufmann 1987, Kosaki et al. 1991).

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Deep-sea Fish of the Hawaiian Archipelago-CHAVE AND MUNDY . 1985, Hourigan and Reese have endemic Japanese species as sister taxa.
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