Abstract
The dark-mouth skate, Raja arctowskii Dollo, 1904 from Antarctic waters is an extraordinary case in skate taxonomy. For more than 100 years, this species has been known only from three empty egg capsules and the species as such has remained undescribed due to the lack of specimens that could be assigned to Dollo’s small capsules. Since trawled egg capsules and an egg capsule containing a near-term embryo became available, it finally was possible to connect specimens with the empty egg capsules and completely describe Dollo’s R. arctowskii with detailed external morphology, skeletal features, clasper morphology, and clasper skeleton and assign it to the genus Bathyraja Ishiyama, 1958a. Bathyraja arctowskii is one of the smallest known species of Bathyraja, attaining only a 61 cm total length (TL). It is characterized by an at least partly, usually completely medium to dark grayish pigmented mouth cavity, as well as the often dark underside of the nasal curtain from very small juvenile stages onwards. It further differs from most congeners in Antarctic and Subantarctic waters in the absence of thorns on the dorsal disc. It appears to be a wide-ranging, circumantarctic species found in the Atlantic, Pacific, and Indian Ocean sectors of the Southern Ocean. The species seems to be locally common at least in the Atlantic sector, with up to 94 juvenile to subadult specimens caught in one single haul.
Similar content being viewed by others
References
Bean TH (1881) Descriptions of new fishes from Alaska and Siberia. Proc U S Natl Mus 4(210):144–159
Beebe W, Tee-Van J (1941) Eastern Pacific expeditions of the New York Zoological Society. XXVIII. Fishes from the Tropical Eastern Pacific. [From Cedros Island, Lower California, South to the Galápagos Islands and Northern Peru.] Part 3. Rays, mantas and chimaeras. Zool Sci Contrib NY Zool Soc 26:245–280
Bigelow HB, Schroeder WC (1953) Fishes of the Western North Atlantic. Part two. Sawfishes, guitarfishes, skates and rays, Chimaeroids. Mem Sears Found Mar Res 1(2):1–588
Bigelow HB, Schroeder WC (1965) Notes on a small collection of rajids from the sub-Antarctic region. Limnol Oceanogr 10(Suppl):R38–R49
Clark RS (1926) Rays and skates, a revision of the European species. Fish Bd Scotland Sci Inv 1:1–66
Concha F, Hernández S, Oddone MC (2009) Egg capsules of the raspthorn sandskate, Psammobatis scobina (Philippi, 1857) (Rajiformes, Rajidae). Rev Biol Mar Oceanogr 44(1):253–256
Concha F, Oddone MC, Bustamante C, Morales N (2012) Egg capsules of the yellownose skate Zearaja chilensis (Guichenot 1848) and the roughskin skate Dipturus trachyderma (Krefft and Stehmann 1974) (Rajiformes: Rajidae) from the South-Eastern Pacific Ocean. Ichthyol Res 59:323–327
de Buen F (1959) Notas preliminares sobre la fauna marina preabismal de Chile, con descripción de una familia de rayas, dos géneros y siete especies nuevos. Bol Mus Nac Chile 27(3):171–201
Díaz de Astarloa JM, Mabragaña E (2004)Bathyraja cousseauae sp. n.: a new softnose skate from the southwestern Atlantic (Rajiformes, Rajidae). Copeia 2004(2):326–335
Dolganov VN (1983) Rukovodstvo po opredeleniyu khryashchevykh ryb dal’nevostochnykh morei SSSR i sopredel’nykh vod [Manual for identification of cartilaginous fishes of Far East seas of USSR and adjacent waters]. TINRO, Vladivostok
Dolganov VN (1985) A new species of skate from the southwestern Pacific Ocean. Biol Morya (Vladivostok) 1985(5):66–68
Dollo L (1904) Poissons. In: Expédition Antarctique Belge, Résultats du voyage du S.Y. Belgica en 1897-1898-1899, Zoologie. Imprimerie J.-E. Buschmann, Antwerpen, pp. 1–240
Duhamel G, Gasco N, Davaine P (2005) Poissons des îles Kerguelen et Crozet. Guide régional de l’océan Austral. Muséum national d’Histoire naturelle, Paris
Ebert DA (2016) Deep–sea cartilaginous fishes of the southeastern Pacific Ocean. FAO species catalogue for fishery purposes No. 10. FAO, Rome
Ebert DA, Davis CD (2007) Descriptions of skate egg cases (Chondrichthyes: Rajiformes: Rajoidei) from the eastern North Pacific. Zootaxa 1393:1–18
Forster GR (1967) A new deep-sea ray from the Bay of Biscay. J Mar Biol Assoc U K 47(2):281–286
Fowler HW (1910) Notes on batoid fishes. Proc Acad Nat Sci Phila 62:468–475
Garrick JAF (1961) Studies on New Zealand Elasmobranchii. Part XIII. A new species of Raja from 1,300 Fathoms. Trans R Soc N Z 88(4):743–748
Gilbert CH (1892) Descriptions of thirty-four new species of fishes collected in 1888 and 1889, principally among the Santa Barbara Islands and in the Gulf of California. In: scientific results of explorations by the U. S Fish Commission steamer Albatross. Proc U S Natl Mus 14(880):539–566
Gilbert CH (1896) The ichthyological collections of the steamer Albatross during the years 1890 and 1891. Rep U S Fish Comm 19:393–476
Gill TN, Townsend CH (1897) Diagnoses of new species of fishes found in Bering Sea. Proc Biol Soc Wash 11:231–234
Günther A (1876) Remarks on fishes, with descriptions of new species in the British Museum, chiefly from southern seas. Ann Mag Nat Hist (Series 4) 17(101):389–402
Günther A (1877) Preliminary notes on new fishes collected in Japan during the expedition of H.M.S. ‘Challenger’. Ann Mag Nat Hist (Series 4) 20(119):433–446
Günther A (1880) Report on the shore fishes procured during the voyage of H.M.S. Challenger in the years 1873–1876. In: Report on the scientific results of the voyage of H.M.S. Challenger during the years 1873–76. Zoology 1(part 6):1–82
Hanchet SM, Stewart AL, McMillan PJ, Clark MR, O’Driscoll RL, Stevenson ML (2013) Diversity, relative abundance, new locality records, and updated fish fauna of the Ross Sea region. Antarct Sci 25(5):619–636
Hubbs CL, Ishiyama R (1968) Methods for the taxonomic study and description of skates (Rajidae). Copeia 1968(3):483–491
Hulley PA (1970) An investigation of the Rajidae of the west and south coasts of southern Africa. Ann S Afr Mus 55(4):151–220
Hulley PA (1972) The origin, interrelationships and distribution of southern African Rajidae (Chondrichthyes, Batoidei). Ann S Afr Mus 60:1–103
Hureau J-C, Ozouf-Costaz C (1980) Une nouvelle espèce de raie bathyale des îles Kerguelen Bathyraja irrasa n. sp. (Chondrichthyes, Rajidae). Cybium 4(9):19–29
Iglésias SP, Lévy-Hartmann L (2012)Bathyraja leucomelanos, a new species of softnose skate (Chondrichthyes: Arhynchobatidae) from New Caledonia. Ichthyol Res 59(1):38–48
Ishihara H, Treloar M, Bor PHF, Senou H, Jeong CH (2012) The comparative morphology of skate egg capsules (Chondrichthyes: Elasmobranchii: Rajiformes). Bull Kanagawa Prefect Mus (Nat Sci) 41:9–25
Ishiyama R (1952) Studies on the rays and skates belonging to the family Rajidae, found in Japan and adjacent regions. 4. A revision of three genera of Japanese rajids with descriptions of one new genus and four new species mostly occured in northern Japan. J Shimonoseki Coll Fish 2(2):1–34
Ishiyama R (1958a) Studies on the rajid fishes (Rajidae) found in the waters around Japan. J Shimonoseki Coll Fish 7(2/3):193–394
Ishiyama R (1958b) Observation of the egg-capsules of skates of the family Rajidae, found in Japan and adjacent waters. Bull Mus Comp Zool 118(1):1–24
Ishiyama R (1967) Fauna japonica. Rajidae (Pisces). Biogeographical Society of Japan, Tokyo
Ishiyama R, Ishihara H (1977) Five new species of skates in the genus Bathyraja from the western North Pacific, with reference to their interspecific relationships. Jpn J Ichthyol 24(2):71–90
Jensen AS (1914) The selachians of Greenland. Bianco Lunos Bogtrykkeri, København
Jones C, Damerau M, Deitrich K, Driscoll R, Kock KH, Kuhn K, Moore J, Morgan T, Near T, Pennington J, Schöling S (2009) Chapter 9. Demersal finfish survey of the South Orkney Islands. In: Van Cise AM (ed) AMLR 2008/2009 field season report – objectives, accomplishments and tentative conclusions, NOAA-TM-NMFS-SWFSC-445. U.S. Department of Commerce, National Oceanic & Atmospheric Administration, La Jolla, pp 49–66
Kalisz H (2013) The comparison of two undescribed species of skates, Bathyraja sp. 2 and Bathyraja sp. (c.f. eatonii) from the Antarctic waters of the South Orkney Islands. BSc thesis, Wilkes Honors College of Florida Atlantic University
Kendall WC, Radcliffe L (1912) The shore fishes. Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz by the U.S. fish commission steamer ‘Albatross’ from October, 1904 to March, 1905, Lieut. Commander L.M. Garrett, U.S.N. commanding. Mem Mus Comp Zool 35(3):75–171
Krefft G (1968a) Knorpelfische (Chondrichthyes) aus dem tropischen Ostatlantik. Atlantide Rep 10:33–76
Krefft G (1968b) Neue und erstmalig nachgewiesene Knorpelfische aus dem Archibenthal des Südwestatlantiks, einschließlich einer Diskussion einiger Etmopterus-Arten südlicher Meere. Arch Fischereiwiss 19(1):1–42
Last PR, Séret B (2008) Three new legskates of the genus Sinobatis (Rajoidei: Anacanthobatidae) from the Indo-West Pacific. Zootaxa 1671:33–58
Last PR, Weigmann S, Yang L (2016a) Changes to the nomenclature of the skates (Chondrichthyes: Rajiformes). In: Last PR, Yearsley GK (eds) Rays of the world: supplementary information. CSIRO Special Publication. CSIRO Publishing, Melbourne, pp. 11–34
Last PR, Stehmann MFW, Séret B, Weigmann S (2016b) Softnose skates, family Arhynchobatidae. In: Last PR, White WT, de Carvalho MR, Séret B, Stehmann MFW, Naylor GJP (eds) Rays of the world. CSIRO Publishing, Melbourne, pp 364–472
Last PR, Stewart AL, Séret B (2016c) A new temperate deepwater skate of the genus Bathyraja (Rajoidei: Arhynchobatidae) from the South-West Pacific. Zootaxa 4132(1):107–117
Matsuoka K, Skoglund A, Roth G (2018) Quantarctica [Dataset]. Nor Polar Inst. https://doi.org/10.21334/npolar.2018.8516e961
McEachran JD (1983) Results of the research cruises of FRV “Walther Herwig” to South America. LXI. Revision of the South American skate genus Psammobatis Günther, 1870 (Elasmobranchii: Rajiformes, Rajidae). Arch Fischereiwiss 34(1):23–80
McEachran JD, Compagno LJV (1979) A further description of Gurgesiella furvescens with comments on the interrelationships of Gurgesiellidae and Pseudorajidae (Pisces, Rajoidei). Bull Mar Sci 29(4):530–553
McEachran JD, Miyake T (1984) Comments on the skates of the tropical eastern Pacific: one new species and three new records (Elasmobranchii: Rajiformes). Proc Biol Soc Wash 97(4):773–787
Müller J, Henle FGJ (1838–41) Systematische Beschreibung der Plagiostomen. Veit und Comp., Berlin
Naylor GJP, Ryburn JA, Fedrigo O, Lopez A (2005) Phylogenetic relationships among the major lineages of modern elasmobranchs. In: Hamlett WC (ed) Reproductive biology and phylogeny of chondrichthyes: sharks, rays and chimaeras, vol 3. Science Publishers, Endfield, pp 1–25
Naylor GJP, Caira JN, Jensen K, Rosana KAM, White WT, Last PR (2012) A DNA sequence-based approach to the identification of shark and ray species and its implications for global elasmobranch diversity and parasitology. Bull Am Mus Nat Hist 367:1–263
Norman JR (1937) Coast fishes. Part II. The Patagonian region. In: Discovery reports Vol. 16. University Press, Cambridge, pp. 1–150
Orr JW, Stevenson DE, Hoff GR, Spies IB, McEachran JD (2011) Bathyraja panthera, a new species of skate (Rajidae: Arhynchobatinae) from the western Aleutian Islands, and resurrection of the subgenus Arctoraja Ishiyama. NOAA Prof Pap NMFS 11:1–50
Osburn RC, Nichols JT (1916) Shore fishes collected by the “Albatross” expedition in Lower California, with descriptions of new species. Bull Am Mus Nat Hist 35:139–181
Philippi RA (1902) Descripcion de cinco nuevas especies chilenas: del órden de los plagióstomos. Anal Univ Chile 109:303–315
Porcu C, Marongiu MF, Bellodi A, Cannas R, Cau A, Melis R, Mulas A, Soldovilla G, Vacca L, Follesa MC (2017) Morphological descriptions of the eggcases of skates (Rajidae) from the central-western Mediterranean, with notes on their distribution. Helgol Mar Res 71(10):1–14
Sabaj MH (2019) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference, Version 7.1 (21 March 2019). Available from http://www.asih.org/ (accessed 11 Mar 2020)
Smith PJ, Steinke D, Mcveagh SM, Stewart AL, Struthers CD, Roberts CD (2008) Molecular analysis of Southern Ocean skates (Bathyraja) reveals a new species of Antarctic skate. J Fish Biol 73(5):1170–1182
Soldatov VK, Pavlenko MN (1915) Description of a new species of family Rajidae from Peter the Great Bay and from Okhotsk Sea. Annuaire Mus Zool Acad Imp Sci St Petersb 20:162–163
Springer S (1971) Three species of skates (Rajidae) from the continental waters of Antarctica. In: Llano GA, Wallen IE (eds) Biology of the Antarctic seas IV, vol 17. American Geophysical Union, Washington D.C., pp 1–10
Springer VG, Garrick JAF (1964) A survey of vertebral numbers in sharks. Proc U S Nat Mus 116(3496):73–96
Stehmann M (1970) Vergleichend morphologische und anatomische Untersuchungen zur Neuordnung der Systematik der nordostatlantischen Rajidae (Chondrichthyes, Batoidei). Arch Fischereiwiss 21(2):73–164
Stehmann M (1985) Ergebnisse der Forschungsreisen des FFS “Walther Herwig” nach Südamerika. LXIV. Bathyraja papilionifera sp. n. (Pisces, Batoidea, Rajidae), eine weitere neue Rochenart aus dem Südwestatlantik vom nordargentinischen Kontinentalabhang. Arch Fischereiwiss 36(1/2):195–211
Stehmann M (1987) Bathyraja meridionalis sp. n. (Pisces, Elasmobranchii, Rajidae), a new deep-water skate from the eastern slope of subantarctic South Georgia Island. Arch Fischereiwiss 38(1–2):35–56
Stehmann MFW (1995) First and new records of skates (Chondrichthyes, Rajiformes, Rajidae) from the West African continental slope (Morocco to South Africa), with descriptions of two new species. Arch Fish Mar Res 43(1):1–119
Stehmann MFW (2005a) Bathyraja ishiharai n. sp., a new deep-water skate from the eastern Indian Ocean on the Naturalist Plateau off southwestern Australia (Elasmobranchii, Rajiformes, Rajidae). J Ichthyol 45(suppl. 1):S39–S57
Stehmann MFW (2005b) Bathyraja tunae n. sp., a new deep-water skate from the western Indian Ocean (Chondrichthyes, Rajiformes, Rajidae). J Ichthyol 45(suppl. 1):S23–S38
Stehmann M, Bürkel DL (1990) Rajidae. In: Gon O, Heemstra PC (eds) Fishes of the Southern Ocean. J.L.B. Smith Institute of Ichthyology, Grahamstown, pp 86–97
Stehmann MFW, Séret B, Costa EM, Baro J (2008) Neoraja iberica n. sp., a new species of pygmy skate (Elasmobranchii, Rajidae) from the southern upper slope of the Iberian Peninsula (Eastern North Atlantic). Cybium 32(1):51–71
Stevenson DE, Orr JW, Hoff GR, McEachran JD (2004)Bathyraja mariposa: a new species of skate (Rajidae: Arhynchobatinae) from the Aleutian Islands. Copeia 2004(2):305–314
Suvorov EK (1935)Raja violacea, a new species of ray from the Okhotsk Sea [in Russian with English summary]. Bull Acad Sci Math Nat 1935:431–433
Tanaka S (1916) Four new species of Japanese fishes [in Japanese]. Dobutsugaku Zasshi 28(331):173–174
Treloar MA, Laurenson LJB, Stevens JD (2006) Descriptions of rajid egg cases from southeastern Australian waters. Zootaxa 1231:53–68
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(3):837–1037
Weigmann S (2017) Reply to Borsa (2017): comment on ‘annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity by Weigmann (2016)’. J Fish Biol 90(4):1176–1181
Acknowledgments
The authors are very grateful to Bernard Séret (ICHTYO CONSULT) for morphometrics, meristics, and description notes of the MNHN specimen and Jonathan Pfliger, Guy Duhamel, and Zouhaira Gabsi (MNHN) for photographs and radiographs of this specimen, as well as basic measurements and photographs of several egg capsules of B. murrayi. Guy Duhamel also kindly granted permission to use his fresh photograph of the MNHN specimen in the present publication. The authors are also very grateful to Olivier Pauwels (IRSNB) for kindly granting permission to use photographs of the syntype egg capsule IRSNB 25 in the present publication. Lei Yang (FLMNH) generated the molecular data used in the study. The authors further would like to thank Hajime Ishihara (W&I Associates Co. Ltd., Japan) for providing an image of an egg capsule and embryo of B. murrayi and granting permission to use this photograph in the present publication; Andrew Stewart, and Carl Struthers (NMNZ) for detailed data, photographs, and tissue samples of specimens at NMNZ; Will White (CSIRO) for sending tissue samples of Bathyraja cf. eatonii, B. eatonii, B. irrasa, B. maccaini, and B. murrayi; and Jürgen Pollerspöck (Shark-References) for providing references. The second author is also very grateful to Ralf Thiel (ZMH) for granting access to and loan of specimens in the ZMH collection for examinations and access to photo- and radiography facilities at ZMH, as well as for approving dissections of disintegrated specimens, and Irina Eidus (ZMH) for her help with the radiography, collection database and cards, as well as for providing photographs of several specimens. The authors are grateful to journal editors and reviewers of the manuscript for their helpful comments.
Funding
This work was supported (in part) by the NZ National Institute of Water and Atmospheric Research Ltd. Core Funded Coasts & Oceans Programme 2: Biological Resources subcontract for fundamental knowledge of marine fish biodiversity with the Museum of New Zealand Te Papa Tongarewa. The collection of sequence data was supported with an award from the U.S. NSF Foundation to GJPN, DEB 01132229.
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Conflict of interest
The authors declare that they have no conflict of interest.
Ethical approval
All applicable international, national, and/or institutional guidelines for animal testing, animal care, and use of animals were followed by the authors.
Sampling and field studies
All necessary permits for sampling and observational field studies have been obtained by the authors from the competent authorities and are mentioned in the acknowledgements, if applicable. The study is compliant with CBD and Nagoya protocols.
Data availability
All data generated or analyzed during this study are included in this published article.
Author contribution
MFWS and SW conceived and designed research, conducted examinations, and wrote the first draft of the manuscript, GJPN conducted molecular analyses and contributed parts of the manuscript. All authors read and approved the final manuscript.
Additional information
Communicated by D. Ebert
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
This article is a contribution to the Topical Collection Systematics and Biodiversity of Indian Ocean Sharks, Rays, and Chimaeras (Chondrichthyes)
Rights and permissions
About this article
Cite this article
Stehmann, M.F.W., Weigmann, S. & Naylor, G.J.P. First complete description of the dark-mouth skate Raja arctowskii Dollo, 1904 from Antarctic waters, assigned to the genus Bathyraja (Elasmobranchii, Rajiformes, Arhynchobatidae). Mar. Biodivers. 51, 18 (2021). https://doi.org/10.1007/s12526-020-01124-1
Received:
Revised:
Accepted:
Published:
DOI: https://doi.org/10.1007/s12526-020-01124-1