A preliminary review and in situ observations of the spookfish genus Harriotta (Holocephali: Rhinochimaeridae)


The taxonomic status of two species of the spookfish genus Harriotta Goode and Bean 1895 is revised and discussed. On the basis of a literature review along with a comparison between the two species formerly considered valid, H. haeckeli Karrer 1972 is synonymized with H. raleighana Goode and Bean 1895, and Anteliochimaera chaetirhampha Tanaka 1909 previously regarded as a junior synonym of the latter is resurrected as a valid species. The revised H. raleighana is newly recorded from Japan as well as the North Pacific based on two voucher specimens collected from Hyuga-nada off Kyushu, southern Japan; the record extends the species’ known range approximately 8000 km northward from the South Pacific. In addition, secondary sexual characteristics of H. raleighana are documented for the first time, which further illuminates pronounced differences between the two species of Harriotta; H. raleighana is also revealed to have a smaller size at maturity compared with H. chaetirhampha. In the northwestern Pacific, the species show bathymetric zonation with a narrow sympatric zone at depths around 1600 m (1535–2611 m in H. raleighana vs. 906–1654 m in H. chaetirhampha). In situ behavior of these species is described and discussed from video footage.

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  1. Bean BA, Weed AC (1910) Notes on Anteliochimaera and related genera of chimaeroid fishes. Proc US Nat Mus 17:661–663, pl 38

  2. Compagno LJV, Stehmann M, Ebert DA (1990) Rhinochimaera africana, a new longnose chimaera from southern Africa, with comments on the systematics and distribution of the genus Rhinochimaera Garman, 1901 (Chondrichthyes, Chimaeriformes, Rhinochimaeridae). S Afr J Mar Sci 9:201–222

  3. Deng S-M, Xiong G-Q, Zhan H-X (1983) Description of three new species of elasmobranchiate fishes from deep waters of the East China Sea. Oceanol Limnol Sin 14:64–70

  4. Didier DA, Nakaya K (1999) Redescription of Rhinochimaera pacifica (Mitsukuri) and first record of R. africana Compagno, Stehmann & Ebert from Japan (Chimaeriformes: Rhinochimaeridae). Ichthyol Res 46:139–152

  5. Didier DA, Stehmann M (1996) Neoharriotta pumila, a new species of longnose chimaera from the northwestern Indian Ocean (Pisces, Holocephali, Rhinochimaeridae). Copeia 1996:955–965

  6. Didier DA, Kemper JM, Ebert DA (2012) Phylogeny, biology, and classification of extant holocephalans. In: Carrier JC, Musick JA, Heithaus MR (eds) Biology of sharks and their relatives, 2nd edn. CRC Press, Boca Raton, pp 97–122

  7. Ebert DA (2014) Deep-sea cartilaginous fishes of the Indian Ocean, vol 2. Batoids and chimaeras. FAO, Rome

  8. Ebert DA, Mostarda E (2013) Identification guide to the deep-sea cartilaginous fishes of the Indian Ocean. FAO, Rome

  9. Finucci B, Dunn MR, Jones EG, Anderson J (2017) Reproductive biology of the two deep-sea chimaerids, longnose spookfish (Harriotta raleighana) and Pacific spookfish (Rhinochimaera pacifica). Deep-Sea Res I 120:76–87

  10. Fowler HW (1941) Contributions to the biology of the Philippine Archipelago and adjacent regions. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostraophysi obtained by the United States Bureau of fisheries steamer “Albatross” in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. US Nat Mus Bull 13: i–x + 1–879

  11. Fricke R, Eschmeyer WN (2019) A guide to fish collections in the Catalog of Fishes. Accessed 23 January 2019

  12. Fricke R, Eschmeyer WN, van der Laan R (2019) Catalog of Fishes: genera, species, references. Accessed 23 January 2019

  13. Goode GB, Bean TH (1895) On Harriotta, a new type of chimaeroid fish from the deep waters of the northwestern Atlantic. Proc US Nat Mus 17:471–473, pl XIX

  14. Gosline WA (1971) Functional morphology and classification of teleostean fishes. University of Hawaii Press, Honolulu

  15. Hoving HJT, Robison BH (2016) Deep-sea in situ observations of gonatid squid and their prey reveal high occurrence of cannibalism. Deep-Sea Res I 116:94–98

  16. ICZN (International Commission on Zoological Nomenclature) (1999) International code of zoological nomenclature. 4th edn. International Trust for Zoological Nomenclature, London

  17. JAMSTEC (Japan Agency for Marine-Earth Science and Technology) (2019) JAMSTEC E-library of Deep-sea Images (J-EDI). Accessed 23 January 2019

  18. Jordan DS, Evermann BW (1900) The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part IV. Bull US Nat Mus 47:i–ci + 3137–3313, pls I–CCCXCII

  19. Jordan DS, Tanaka S, Snyder JO (1913) A catalogue of the fishes of Japan. J Coll Sci Tokyo Imp Univ 33:1–497

  20. Karrer C (1972) Die gattung Harriotta Goode and Bean, 1985 (Chondrichthyes, Chimaeriformes, Rhinochimaeridae). Mitt Zool Mus Berl 48:203–221

  21. Last PR, Stevens JD (2009) Sharks and rays of Australia, 2nd edn. CSIRO Publishing, Clayton

  22. Lauder GV (2000) Function of the caudal fin during locomotion in fishes: kinematics, flow visualization, and evolutionary patterns. Am Zool 40:101–122

  23. Lombardi J (1998) Comparative vertebrate reproduction. Kluwer Academic Publishers, Norwell

  24. Nakabo T, Yagishita N, Yamaguchi A (2013) Rhinochimaeridae. In: Nakabo T (ed) Fishes of Japan with pictorial keys to the species, 3rd edn. Tokai University Press, Hadano, pp 148–149, 1755–1756

  25. Nakaya K (1982) Rhinochimaeridae. In: Okamura O, Kitajima T (eds) Fishes of the Okinawa Trough and the adjacent waters I. Japan Fisheries Resource Conservation Association, Tokyo, pp 76–77, 313

  26. Nakaya K (1988) Rhinochimaeridae. In: Masuda H, Amaoka K, Araga C, Uyeno T, Yoshino T (eds) The fishes of the Japanese Archipelago, 2nd edn. Tokai University Press, Tokyo, p 17, pls 20, 337

  27. Patterson C (1968) The caudal skeleton in Lower Liassic pholidophorid fishes. Bull Br Mus Nat Hist Geol 16:203–239

  28. R Development Core Team (2015) R: A Language and environment for statistical computing. R Foundation for Statistical Computing, Vienna

  29. Senou H (2012) Chapter 31. Vertebrate. In: Fujikura K, Okutani T, Maruyama T (eds) Deep-sea life—Biological observations using research submersibles, 2nd edn. Tokai University Press, Hadano, pp 356–382

  30. Stehmann M, Bürkel DL (1984) Rhinochimaeridae. In: Whitehead PJP, Bauchot M-L, Hureau J-C, Nielsen J, Tortonese E (eds) Fishes of the north-eastern Atlantic and the Mediterranean. Vol I. UNESCO, Paris, pp 216–218

  31. Tanaka S (1909) Descriptions of one new genus and ten new species of Japanese fishes. J Coll Sci Imp Univ Tokyo 27 (art 8):1–27, pl I

  32. Townsend CH, Nichols JT (1925) Deep sea fishes of the ‘Albatross’ Lower California Expedition. Bull Am Mus Nat Hist 52:1–20, pls I–IV

  33. Weigmann S (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. J Fish Biol 88:837–1037

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We are grateful to D. A. Didier (Millersville University, Millersville) for sharing her knowledge on rhinochimaerid taxonomy, S. Tanaka (Tokai University, Shizuoka) for providing valuable comments on this study, and K. Sakamoto (ZMUT) for his curatorial assistance; K. Murasaki (Tokai University, Shizuoka) kindly provided photographs of the lectotype of H. raleighana. We also thank Y. Toyoda (JAMSTEC, Yokohama) for helping us to access the high-resolution footage examined here, and G. Yearsley (Hobart) for editing the English text. This study was supported by JSPS KAKENHI Grant Number JP18K14509.

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Correspondence to Naohide Nakayama.

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Nakayama, N., Matsunuma, M. & Endo, H. A preliminary review and in situ observations of the spookfish genus Harriotta (Holocephali: Rhinochimaeridae). Ichthyol Res 67, 82–91 (2020) doi:10.1007/s10228-019-00703-y

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  • Chimaeriformes
  • Synonymy
  • Range extension
  • Secondary sexual characteristics
  • Bathymetric zonation
  • Japan