Pallas, 1814) and a Redescription of Sebastes Ciliatus (Tilesius, 1813
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The dusky rockfishes (Teleostei: Socrpaeniformes) of the North Pacific Ocean: resurrection of Sebastes variabilis (Pallas, 1814) and a redescription of Sebastes ciliatus (Tilesius, 1813) Item Type article Authors Orr, James W.; Blackburn, James E. Download date 26/09/2021 03:53:40 Link to Item http://hdl.handle.net/1834/30907 328 Abstract— The dusky rockfish (Se- The dusky rockfishes (Teleostei: Scorpaeniformes) bastes ciliatus) of the North Pacific Ocean has been considered a single of the North Pacific Ocean: resurrection of variable species with light and dark forms distributed in deep and shal- Sebastes variabilis (Pallas, 1814) and a low water, respectively. These forms redescription of Sebastes ciliatus (Tilesius, 1813) have been subjected to two distinct fisheries separately managed by fed- eral and state agencies: the light deep James W. Orr form is captured in the offshore trawl Resource Assessment and Conservation Engineering Division fishery; the dark shallow form, in the Alaska Fisheries Science Center nearshore jig fishery. The forms have National Marine Fisheries Service, NOAA been commonly recognized as the light 7600 Sand Point Way NE dusky and dark dusky rockfishes. From Seattle, Washington 98115 morphological evidence correlated with E-mail address: [email protected] color differences in some 400 speci- mens, we recognize two species cor- responding with these color forms. James E. Blackburn Sebastes ciliatus (Tilesius) is the dark Alaska Department of Fish and Game shallow-water species found in depths 211 Mission Road of 5−160 m in the western Aleutian Kodiak, Alaska 99615-9988 Islands and eastern Bering Sea to British Columbia. The name Sebastes variabilis (Pallas) is resurrected from the synonymy of S. ciliatus to apply to the deeper water species known from depths of 12−675 m and ranging from Among the approximately 92 species agencies since 1998. The light-colored Hokkaido, Japan, through the Aleu- of Sebastes found in the North Pacific, deep form is captured in the offshore tian Islands and eastern Bering Sea, to Oregon. Sebastes ciliatus is uniformly two commercially important species trawl fishery and is the dominant spe- dark blue to black, gradually lightening long identified under the name Sebastes cies of the pelagic shelf rockfish fisher- on the ventrum, with a jet black peri- ciliatus have been taxonomically prob- ies complex regulated by the National toneum, a smaller symphyseal knob, lematic. The name S. ciliatus (Tilesius, Marine Fisheries Service (NMFS). and fewer lateral-line pores compared 1813) has been commonly applied to Specific catch limits are set under the to S. variabilis. Sebastes variabilis is specimens considered to represent a designation “dusky rockfish.” The oc- more variable in body color, ranging single variable species ranging from casional catch of the dark form in these from light yellow to a more usual tan northern Japan to British Columbia offshore waters has also been consid- or greenish brown to a nearly uniform (Barsukov, 1964; Westrheim, 1973; ered “dusky rockfish.” The dark-colored dark dorsum, but it invariably has a Shinohara et al., 1994; Mecklenburg et shallow form is found commonly in the distinct red to white ventrum. Syn- onymies, diagnoses, descriptions, and al., 2002), and the name S. variabilis nearshore jig fishery regulated by the geographic distributions are provided (Pallas, 1814) has been treated as a Alaska Department of Fish and Game. for each species. junior synonym (Jordan and Gilbert, The dark form, routinely misidentified 1881; Eigenmann and Beeson, 1894; as S. melanops, may comprise up to 25% Jordan and Evermann, 1898; Blanc of the catch in the “black rockfish” jig and Hureau, 1968). Two color forms fishery off Kenai Peninsula (Clausen within S. ciliatus have been reported et al.1) and is managed only as “other and hypothesized to be distinct species rockfish” bycatch within the fishery. (Quast and Hall, 1972; Eschmeyer et Early allozyme analyses (Tsuyuki al., 1983; Kessler, 1985; Fig. 1). The et al., 1965, 1968) indicated signifi- typically light-colored form, commonly cant genetic differences among samples known as the light dusky rockfish, is identified as S. ciliatus. A more detailed often found in large aggregations over analysis of several Sebastes species the outer continental shelf and upper (Seeb, 1986) and a later study focused slope at depths down to 675 m, and less frequently in nearshore habitats. The dark-blue to black form, commonly known as the dark dusky rockfish, is 1 Clausen, D. M., C. R. Lunsford, and found in more shallow habitats from J. T. Fujioka. 2002. Pelagic shelf Manuscript approved for publication nearshore rocky reefs to depths no rockfish. In Stock assessment and fish- 22 December 2003 by Scientific Editor. greater than 160 m. ery evaluation report for the groundfish resources of the Gulf of Alaska for 2002, p. Manuscript received 20 January 2004 These forms have been subjected to 383–417. North Pacific Fishery Manage- at NMFS Scientific Publications Office. two distinct fisheries separately man- ment Council, 605 W 4th Ave, Suite 306, Fish. Bull. 102:328–348 (2004). aged by U.S. federal and Alaska state Anchorage, AK 99501. Orr and Blackburn: Resurrection of Sebastes variabilis and redescription of Sebastes ciliatus 329 A B C D E Figure 1 (A) Sebastes variabilis (top), UW 43494, 225.2 mm, and S. ciliatus (bottom), UW 43493, neotype, 266.4 mm, collected at 37 and 25 m depth, respectively, in Lynn Canal near Funter Bay, southeast Alaska. (B) Sebastes ciliatus (top), UW 45512, 235.2 mm, and S. variabilis (bottom), UW 45511, 206.2 mm, collected syntopically at 67 m depth in the northern Gulf of Alaska, 57.38061°N, 154.8009°W. (C) Sebastes variabilis (left), UW 43494, 150.6−225.2 mm, and S. ciliatus (right), UW 43492, 153.7−241.1 mm, collected at 37 and 25 m depth, respectively, in Lynn Canal near Funter Bay, southeast Alaska. (D) Sebastes variabilis, UW 43251, 390 mm (top) and 410 mm (bottom), northern Gulf of Alaska, 59.50446°N, 145.2262°W, 135 m depth. (E) Sebastes melanops (top), UW 43490, and S. ciliatus (bottom), UW 43484, 313 mm, collected at 25 m depth in Soapstone Cove, southeast Alaska. 330 Fishery Bulletin 102(2) A B Figure 2 (A) Perca variabilis Pallas, MNHN 8670, lectotype, 343.7 mm, “mari Americam borealum.” (B) Epinephelus ciliatus Tilesius, illustration of holotype after Tilesius (1813), specimen presumed lost. on S. ciliatus (Westrheim and Seeb2) concluded that the from NADH dehydrogenase subunit regions of the mito- two color forms were distinct sister species. Seeb’s recent chondrial genome, however, have not revealed differences work with microsatellite DNA data has revealed discrete between the two forms (López4, Gray5), nor have sequence genetic differences between the two, as well as some evi- data from other work on closely related species of Sebastes dence for infrequent hybridization (Seeb3). Sequence data (Bentzen et al., 1998; Sundt and Johansen, 1998; Roques et al., 2001). 2 Westrheim, S. J., and L. W. Seeb. 1997. Unpubl. manu- script. Investigation of the Sebastes ciliatus species group, 36 p. Fisheries and Oceans Canada, Pacific Biological Sta- 4 López, J. A. 2000. Personal commun. Iowa State Univ., tion, Nanaimo, BC, Canada V9R 5K6. Ames, IA 50014. 3 Seeb, L. W. 2002. Personal commun. Alaska Department 5 Gray, A. 2000. Personal commun. Fisheries Division, of Fish and Game, 333 Raspberry Road, Anchorage, AK 99518- School of Fisheries and Ocean Sciences, Univ. Alaska Fair- 1599. banks, 11120 Glacier Highway, Juneau, AK 99801. Orr and Blackburn: Resurrection of Sebastes variabilis and redescription of Sebastes ciliatus 331 In this study, we provide morphological evidence from (Mathsoft, Inc., Seattle, WA). Differences were considered examination of about 400 specimens collected throughout significant at P < 0.05. the geographic and bathymetric range of the species to Arcsine-transformed morphometric ratios (with SL or correlate color differences with meristic and shape differ- head length as denominator) and meristic characters were ences. In thus recognizing two species, S. ciliatus and S. tested to meet the assumptions of normality required for variabilis, previously referred to the name S. ciliatus (Tile- ANOVA. The following characters exhibited normal distri- sius, 1813), we discuss the nomenclatural consequences of butions and did not differ significantly in variance between this decision. Both species were originally described (as species and were subjected to ANOVA: head length, orbit Epinephelus ciliatus Tilesius and Perca variabilis Pallas) length, snout length, interorbital width, suborbital depth, on the basis of early Russian collections from along the gill-raker length, body thickness, pectoral-fin base width, Aleutian Islands (Svetovidov, 1978, 1981). The type series pectoral-fin ray length, caudal peduncle ventral length, of one species is now represented by a single extant speci- predorsal length, spinous dorsal-fin base, soft dorsal-fin men (Fig. 2A) and the other by the illustration of a single, base, and counts of lateral-line pores and gill rakers. now lost, specimen (Svetovidov, 1978, 1981; Fig. 2B). For morphometric characters, significant differences Although workers since the turn of the century have as- were also identified by using an analysis of covariance sociated the name S. ciliatus with the variably light-col- (ANCOVA) of log-10-transformed measurements with SL ored species (Jordan, 1896; Jordan and Evermann, 1898; or head length (HL) as covariates when assumptions of Barsukov, 1964; Orr et al., 1998, 2000; Mecklenburg et normality and the homogeneity of slopes were satisfied. al., 2002), the original description and accompanying il- The ANCOVA model included species as a factor, SL or HL lustration (Fig. 2B) appear to describe the uniformly dark as a covariate, and a species/(SL or HL) interaction (e.g., species. We have also identified the remaining syntype HL=C+Species+SL+(Species×SL)). A residual analysis (Fig. 2A) of Perca variabilis as the light species.