NOAA Professional Paper NMFS 11 U.S. Department of Commerce March 2011 Bathyraja panthera, a new species of skate (Rajidae: Arhynchobatinae) from the western Aleutian Islands, and resurrection of the subgenus James W. Orr Duane E. Stevenson Arctoraja Ishiyama Gerald R. Hoff Ingrid Spies John D. McEachran U.S. Department of Commerce NOAA Professional Gary Locke Secretary of Commerce National Oceanic Papers NMFS and Atmospheric Administration Jane Lubchenco, Ph.D. Scientific Editor Administrator of NOAA Richard D. Brodeur, Ph.D. Associate Editor National Marine Julie Scheurer Fisheries Service Eric C. Schwaab National Marine Fisheries Service Assistant Administrator Northwest Fisheries Science Center for Fisheries 2030 S. Marine Science Dr. Newport, Oregon 97365-5296 Managing Editor Shelley Arenas National Marine Fisheries Service Scientific Publications Office 7600 Sand Point Way NE Seattle, Washington 98115 Editorial Committee Ann C. Matarese, Ph.D. National Marine Fisheries Service James W. Orr, Ph.D. National Marine Fisheries Service Bruce L. Wing, Ph.D. National Marine Fisheries Service The NOAA Professional Paper NMFS (ISSN 1931-4590) series is published by the Scientific Publications Office, National Marine Fisheries Service, The NOAA Professional Paper NMFS series carries peer-reviewed, lengthy original NOAA, 7600 Sand Point Way NE, research reports, taxonomic keys, species synopses, flora and fauna studies, and data-in- Seattle, WA 98115. tensive reports on investigations in fishery science, engineering, and economics. Copies The Secretary of Commerce has determined that the publication of of the NOAA Professional Paper NMFS series are available free in limited numbers to this series is necessary in the transac- government agencies, both federal and state. They are also available in exchange for tion of the public business required by other scientific and technical publications in the marine sciences. Professional Papers law of this Department. Use of funds are published online in PDF format at http://spo.nmfs.noaa.gov for printing of this series has been ap- proved by the Director of the Office NOTICE: This series was established in 2003 to replace the NOAA Technical Report of Management and Budget. NMFS series. NOAA Professional Paper NMFS 11 Bathyraja panthera, a new species of skate (Rajidae: Arhynchobatinae) from the western Aleutian Islands, and resurrection of the subgenus Arctoraja Ishiyama James W. Orr Duane E. Stevenson Gerald R. Hoff Ingrid Spies John D. McEachran March 2011 U.S. Department of Commerce Seattle, Washington Suggested reference Orr, James W., Duane E. Stevenson, Gerald R. Hoff, Ingrid Spies, and John D. McEachran. 2011. Bathyraja panthera, a new species of skate (Rajidae: Arhyncho- batinae) from the western Aleutian Islands, and resurrection of the subgenus Arctoraja Ishiyama. NOAA Professional Paper NMFS 11, 50 p. Online dissemination This report is posted online in PDF format at http://spo.nmfs.noaa.gov (click on Professional Papers link). Purchasing additional copies Additional copies of this report may be available for purchase in paper copy or microfiche from the National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161; 1-800-553-NTIS; http://www.ntis.gov. Copyright law Although the contents of the Professional Papers have not been copyrighted and may be reprinted entirely, reference to source is appreciated. Proprietary products The National Marine Fisheries Service (NMFS) does not approve, recommend, or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication fur- nished by NMFS, in any advertising or sales promotion which would indicate or imply that NMFS approves, recommends, or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. 1 Abstract—We provide morphological Bathyraja panthera, a new species and molecular evidence to recognize a new species of skate from the North Pa- of skate (Rajidae: Arhynchobatinae) cific, Bathyraja panthera. We also resurrect the skate subgenus Arctoraja Ishiyama, from the western Aleutian Islands, confirming its monophyly and the valid- and resurrection of the subgenus Arctoraja Ishiyama ity of the subgenus. Arctoraja was previ- ously recognized as a distinct subgenus of Breviraja and later synonymized with James W. Orr (contact author)1 Bathyraja (family Rajidae). Although the Duane E. Stevenson1 nominal species of Arctoraja have all been 1 considered synonyms of Bathyraja par- Gerald R. Hoff mifera by various authors, on the basis of Ingrid Spies1 morphometric, meristic, chondrological, John D. McEachran2 and molecular data we recognize four 1 species, including the new species. Spe- Resource Assessment and Conservation Engineering Division cies of Arctoraja are distributed across the Alaska Fisheries Science Center North Pacific Ocean and adjacent seas National Marine Fisheries Service, National Oceanic and Atmospheric Administration from southern Japan to British Colum- 7600 Sand Point Way NE bia. Bathyraja parmifera is abundant in Seattle, Washington 98115 the eastern Bering Sea, Aleutian Islands, Email address for contact author: [email protected] and northern Gulf of Alaska; B. smirnovi 2 Department of Wildlife and Fisheries Sciences is a western Pacific species found in the Texas A&M University Sea of Okhotsk and Sea of Japan; B. si- College Station, Texas 77843 moterus is restricted to waters around the northern and eastern coasts of Hokkaido, Japan; and the new species B. panthera is restricted to the western Aleutian Islands. Bathyraja panthera is diagnosed by its color pattern of light yellow blotches with black Introduction at least 25 species are distributed from spotting on a greenish brown background, Japan to California (Ishihara and high thorn and vertebral counts, chondro- Recent bottom-trawl surveys conduct- Ishiyama, 1986; Ishihara, 1990; Ste- logical characters of the neurocranium ed by the Alaska Fisheries Science venson et al., 2004; Ebert and Com- and clasper, and a unique nucleotide sequence within the mitochondrial cyto- Center (AFSC) of the National Ma- pagno, 2007). Twelve species are now chrome oxidase gene. Furthermore, the rine Fisheries Service (NMFS) have considered valid in Alaskan waters species presently recognized as Bathyraja resulted in the collection and subse- (Stevenson et al., 2004; Stevenson and parmifera exhibits two haplotypes among quent discovery of several new species Orr, 2005; Stevenson et al., 2007) and specimens from Alaska, suggesting the and records of fishes from the Aleu- further work is being conducted to possibility of a second, cryptic species. tian Islands, including liparids (Orr clarify the status of suspected species and Busby, 2001, 2006; Orr, 2004; complexes within the region. One of Orr and Maslenikov, 2007), zoarcids these complexes includes the most (Stevenson and Orr, 2006), and skates common species in Alaska, Bathyraja (Stevenson et al., 2004; Stevenson and parmifera, the Alaska skate (Stevenson Orr, 2005). Among those species that et al., 2008) of the subgenus Arctoraja. have been recognized as new but un- Ishiyama (1958a) erected three described is a skate from the western subgenera, Arctoraja, Bathyraja, and Aleutian Islands previously identified Notoraja, for the Japanese skates of as Bathyraja parmifera (Bean, 1881). the widely distributed genus Brevi- Skates of the family Rajidae, domi- raja Bigelow and Schroeder, 1948. nated by the genus Bathyraja, rep- The subgenus Arctoraja included two resent a substantial portion of the species of the North Pacific region, biomass of incidental species encoun- B. parmifera and B. smirnovi (Solda- tered by fisheries in the eastern Ber- tov and Pavlenko, 1915), as well as ing Sea (Spies et al., 2006; Stevenson Ishiyama’s two new subspecies within et al., 2008). Bathyraja Ishiyama, 1958, B. smirnovi: B. s. smirnovi and B. s. comprising over 50 species (Stehm- ankasube. Later, Ishiyama (1967) de- ann, 2005; Ebert and Compagno, scribed Breviraja (Arctoraja) simoterus 2007), is the most diverse genus of based on specimens he had previ- the family, with its major center of ously identified asB. parmifera, yet he abundance in the North Pacific where did not list B. parmifera as a member 2 Professional Paper NMFS 11 of the subgenus. When Ishiyama and Hubbs (1968) elevated the subgenus Bathyraja to generic status, they considered Arctoraja and Notoraja to be synonyms of Bathyraja. Although no comprehensive revision of the sub- genus Arctoraja has been produced since Ishiyama’s (1958a, 1967) seminal works, taxonomic reviews of North Pacific species ofBathyraja have come to contra- dictory conclusions on the status of Bathyraja parmifera and its close relatives. Ishihara and Ishiyama (1985, 1986) treated B. parmifera, B. smirnovi, and B. simoterus as distinct species, as do all recent Japanese authors (Nakaya, 1983, 1984; Nakaya and Shirai, 1992; Nakabo, 2000, 2002), distinguishing them within the context of dichotomous keys and checklists. In their phylogenetic analysis of the Rajidae, McEachran and Dunn (1998) listed all species in a generic classification of species, as did Stehmann (1986), McEachran and Miyake (1990), Figure 1 Compagno (1999), and Ebert and Compagno (2007) Selected terminology used to describe the external in their distributional analyses