355 NOAA First U.S. Commissioner National Marine Fishery Bulletin established 1881 of Fisheries and founder Fisheries Service of Fishery Bulletin Abstract—The diverse predatory A new species of Sebastes (Scorpaeniformes: rockfishes (Sebastes spp.) support extensive commercial fisheries in Sebastidae) from the northeastern Pacific, with the northeastern Pacific. Although 106 species of Sebastes are consid- a redescription of the blue rockfish, S. mystinus ered valid, many of the ecological, (Jordan and Gilbert, 1881) geographical, and morphological boundaries separating them lack clarity. We clarify one such boundary Benjamin W. Frable (contact author)1, 2 by separating the blue rockfish Se- D. Wolfe Wagman2 bastes mystinus (Jordan and Gilbert, Taylor N. Frierson2 1881) into 2 species on the basis of 3 molecular and morphological data. Andres Aguilar We redescribe S. mystinus, desig- Brian L. Sidlauskas1 nate a lectotype, and describe the deacon rockfish, Sebastes diaconus Email address for contact author: [email protected] n. sp. Aside from its unambiguous distinction at 6 microsatellite loci, 1 Department of Fisheries and Wildlife the new species is most easily dif- Oregon State University ferentiated from S. mystinus by its 104 Nash Hall possession of a solid in contrast with Corvallis, Oregon 97331 a blotched color pattern. Sebastes 2 diaconus also possesses a prominent Oregon Department of Fish and Wildlife symphyseal knob versus a reduced Marine Resources Program or absent knob, a flat rather than 2040 SE Marine Science Drive rounded ventrum, and longer first Newport, Oregon 97365 and second anal-fin spines. Sebastes 3 Department of Biological Sciences diaconus occurs from central Cali- California State University, Los Angeles fornia northward to British Colum- 5151 State University Drive bia, Canada, and S. mystinus occurs Los Angeles, California 90032 from northern Oregon south to Baja California Sur, Mexico, indicating a broad region of sympatry in Oregon and northern California. Further collection and study are necessary to clarify distributional boundaries The rockfish genus Sebastes is one of mystinus (Jordan and Gilbert, 1881), and to understand the ecology and the most diverse and abundant gen- black rockfish S. melanops Girard, mechanisms of segregation for this era along the Pacific coast of North 1856, light dusky rockfish S. variabi- species. Additionally, fisheries as- sessments will need revision to ac- America (Love et al., 2002). Rockfish lis (Pallas, 1814), and dusky rockfish count for the longstanding conflation biology and ecology have been well S. ciliatus (Tilesius, 1813), are among of these 2 species. studied because of their commer- the most frequently conflated and cial importance, yet some taxonomic confused when landed in the same limits, population boundaries, and fishery (Kramer and O’Connell, 1995; phylogenetic relationships within Se- Orr and Blackburn, 2004). Even bastes remain unclear (Hyde and Vet- among brightly colored rockfishes, ter, 2007; Orr and Hawkins, 2008) be- increased study has revealed cryptic Manuscript submitted 30 May 2014. cause many species are very similar species. Gharrett et al. (2005) found Manuscript accepted 10 June 2015. and overlap in meristic counts and 2 genetically distinct forms within Fish. Bull. 113:355–377 (2015) morphometrics. As a result, fishery the rougheye rockfish S. aleutianus Online publication date: 9 July 2015. managers struggle to correctly iden- (Jordan and Evermann, 1898), and doi: 10.7755/FB.113.4.1 tify Sebastes species and sometimes those forms were later designated as http://zoobank.org/References/297E1E76- lack accurate species diagnoses to de- the rougheye rockfish S. aleutianus 94C2-49B2-9520-A519B80BC99B termine proper management. and blackspotted rockfish S. mela- The views and opinions expressed or Several clusters of similar spe- nostictus (Matsubara, 1934) (Orr implied in this article are those of the cies within Sebastes merit increased and Hawkins, 2008). The historical author (or authors) and do not necessarily taxonomic attention. For example, concept of the vermilion rockfish S. reflect the position of the National the uniformly dark-colored species of miniatus (Jordan and Gilbert, 1880a) Marine Fisheries Service, NOAA. Sebastes, such as the blue rockfish S. was also shown relatively recently to 356 Fishery Bulletin 113(4) include 2 reproductively isolated entities (Hyde, et al., much earlier than the LGM and that they have subse- 2008). quently expanded ranges to form an area of sympatry In this study, we focus taxonomic attention on S. from central Oregon to northern California. mystinus, a common nearshore species found from Finally, Burford et al. (2011a, 2011b) examined mi- northern Mexico to British Columbia, Canada. In the crosatellite data from 466 type-1 (northern group) and 19th century, S. mystinus was the most commercially 1752 type-2 (southern group) specimens collected from important species in California; now it is mainly tar- Fort Bragg, California, south to Santa Cruz Island, Cali- geted recreationally (Love et al., 2002). Over the last fornia, to determine rates of hybridization (Burford et decade, multiple studies have identified 2 genetically al., 2011a) and year-class compositional and ecological distinct groups of S. mystinus along the Pacific coast differences (Burford et al., 2011b). Burford et al. (2011a) (Cope, 2004; Burford and Larson, 2007; Burford and found no hybridization in northern localities with higher Bernardi, 2008; Burford, 2009; Burford et al., 2011a, co-occurrence, but they identified low levels in South- 2011b), and in several studies the presence of 2 mor- ern California (highest rate of hybridization=4.1%). This phologically unrecognized species have been hypothe- finding, combined with the identification of a Wahlund sized (Burford and Bernardi, 2008; Burford, 2009; Bur- Effect (i.e., lower heterozygosity than expected at ran- ford et al., 2011a, 2011b). dom between the 2 populations), indicates that reproduc- Cope (2004) first identified genetic distinctions be- tive isolation helps maintain the segregation, especially tween northern and southern populations of blue rock- in areas of overlap (Burford et al., 2011a). Burford et al. fish while studying their stock structure. His analysis (2011a, 2011b) considered their results to provide suf- revealed numerous fixed differences in the sequence ficient evidence that the 2 genetic lineages are cryptic of the mitochondrial control region between samples rather than incipient species. from the Oregon–Washington region and samples from Despite the accumulating detail on genetic differen- California waters. These data indicate that the genet- tiation, no study provided a complementary physical ic break occurred in the vicinity of Cape Mendocino, description of the 2 types. Because of the lack of physi- California. cal descriptions or defining characteristics, precise field Burford and colleagues then applied additional mo- identification of Burford’s genetic lineages (type 1 and lecular and phylogeographic analyses to these 2 popula- type 2) has eluded biologists and fishermen alike. tions (Burford and Larson, 2007; Burford and Bernardi, Meanwhile, fisheries biologists acknowledged 2 dif- 2008; Burford, 2009; Burford et al., 2011a, 2011b). In ferent trunk pigmentation patterns in S. mystinus from a combined analysis of mitochondrial (control region) near the Oregon, Washington, and California shores: and nuclear (recombination-activating gene 1 [RAG1]) the “blue-sided” (Fig. 1, A and C) and “blue-blotched” markers and microsatellites for the subgenus Sebasto- rockfishes (Fig. 1, B and D) (Love, 2011), which, as the somus, Burford and Bernardi (2008) were the first to authors of the earlier genetic studies have indicated, propose that the 2 populations might represent differ- match the type-1 and type-2 genetic lineages, respec- ent species. Within S. mystinus, Burford and Bernardi tively (Burford1). That difference in color pattern indi- (2008) identified 2 clades with higher genetic diver- cates that these lineages may be more morphologically gence (FST=0.120) than that found between 2 well- distinguishable than originally thought. established species (Narum et al., 2004), the gopher The wealth of recent genetic work on S. mystinus rockfish Sebastes carnatus (Jordan and Gilbert, 1880b) and the discovery of a color polymorphism that is con- and the black-and-yellow rockfish S. chrysomelas (Jor- gruent with the major genetic break indicate that a dan and Gilbert, 1881), and with much higher genetic formal taxonomic revaluation is overdue. In this study, divergence than that among populations of S. melanops we 1) test whether the morphotypes correspond with (FST=0.032) (Miller et al., 2005). They estimated the di- the types of Burford and Bernardi (2008), 2) character- vergence time of the 2 lineages to be between 780,000 ize the morphological features of the 2 genetic types, 3) and 920,000 years ago, far preceding the Last Glacial use genetic and morphological data to evaluate species Maximum (LGM) and, therefore, refuting Burford and status, and 4) clarify the geographic ranges of the 2 Larsons’s (2007) hypothesis that the LGM caused al- forms. In doing so, we confirm the genetic separation of lopatric speciation within S. mystinus. the 2 color morphs and provide a formal description of Burford and Bernardi (2008) concluded that the 2 the type-1 or blue-sided form as a new species, S. dia- REVIEW genetically distinct groups