An Historical Review of Sebastes Taxonomy and Systematics

ARTHUR W. KENDALL, Jr.

Discovery: The Late 1700’s terested in these fishes for well over a onym of Serranus scriba, and the rock­ The systematics of the speciose genus century. Fernholm and Wheeler (1983) fish was left without a type species for Sebastes1, rockfishes, particularly in the detailed problems associated with the reference. This made way for recogniz­ North Pacific, have challenged ichthy­ early descriptions of Sebastes. The first ing Ascanius (1772) as the author of ologists and others even casually in­ scholarly reference to a rockfish was by Perca norvegica (i.e. Sebastes norvegi­ Linnaeus (1761) who included Norway cus), the most common rockfish of the and Italy as the range of a fish he northeastern Atlantic. 1 Sebastiscus and Hozukius have been consid­ ered subgenera of Sebastes by some and sepa­ described in 1758 and named Perca rate but closely related genera by others. This marina. In his former publication (Lin­ Proliferation of Subgenera: paper will consider each a separate genus and naeus, 1758), he also described Perca 1861–1898 will focus primarily on the taxonomic history of Sebastes. Sebastes is by far the most species­ scriba, a Mediterranean serranid. Reit­ By 1845, similar fish had been de­ rich and widely distributed genus; Sebastiscus erating his description of Perca marina, scribed as species of Sebastes from and Hozukius have three and two species, respec­ tively, and occur only in the northwest Pacific. Linnaeus (1761) cited the common both the eastern and western North Pa­ As will be discussed later, Sebastes and these Norwegian name (uer or rödfisk) for cific. In 1854, Ayres discovered Sebastes other two genera form an unresolved trichotomy the fish from Norway, making it clear paucispinis from off California (Ayres, within the Sebastidae of the suborder Scorpae­ noidei (Ishida, 1994). These three genera could that he was including a rockfish, as well 1854a). Since Sebastes paucispinis is be given the subfamily name Sebastinae, follow­ as a fish from the Mediterranean region very different from Sebastes norvegi­ ing Matsubara (1943). in his description. cus, but clearly related to it, its dis­ Cuvier (1829) was the first to use covery prompted Gill (1861) to erect Arthur W. Kendall, Jr., retired 3 January 2001, the generic name Sebastes, and in his another genus for it: Sebastodes. This was with the Alaska Fisheries Science Center, second, and much more thorough de­ began a trend by Gill and others to National Marine Fisheries Service, NOAA, 7600 scription of the genus (Cuvier and Va­ establish generic or subgeneric group­ Sand Point Way N.E., Seattle, WA 98115. lenciennes, 1829) he included descrip­ ings as more and more similar species tions of species from the North Atlantic were described on both sides of the (Sebastes norvegicus), the Mediterra­ North Pacific during the 1860’s–1880’s ABSTRACT—Following the initial descrip­ nean Sea (S. imperialis = Helicolenus (Fig. 1, 2). By 1898, Jordan and Ev­ tion of a species of Sebastes from the Atlan­ dactylopterus), and the Southern Hemi­ ermann grouped the 55 northeast Pa­ tic in the late 1700’s, in the late 1800’s sphere (S. capensis).2 Cuvier and Va­ cific species that they recognized into the incredible taxonomic diversity of the lenciennes (1829) recognized the con­ 13 subgenera (Jordan and Evermann, genus began to be recognized as more spe­ fusion between the northern fish and 1898). They also mentioned 13 species cies were discovered in northeast Pacific waters. With over 100 species, most of them Perca marina, which he said was from Japan and 3 species from the from the North Pacific, the genus Sebastes “hardly believable”; nevertheless, the Southern Hemisphere. Generally the (rockfishes) now presents taxonomic prob­ name Sebastes marinus (Linnaeus) was characters of a proposed subgenus were lems at every level. For example, although used until recently for this fish from the only those of the type species for the early efforts to understand relationships among the species resulted in the erection North Atlantic. However, Fernholm and subgenus. In most cases the definitions of several subgenera, those and more recent Wheeler (1983) found that the speci­ of the subgenera were inferred rather efforts remain largely unsuccessful. Also, men Linnaeus used as the basis of his than explicitly stated (Appendix I). the position of the genus within the order description was in fact the Mediterra­ Scorpaeniformes, as well as the limits of nean serranid Serranus scriba, so Se­ Confusion: 1863–1900 the genus and the validity of some species are all unresolved. This paper examines bastes (Perca) marina became a syn­ There was considerable disagreement the worldwide history and status of taxo­ among ichthyologists of the late 1800’s nomic studies on Sebastes, and reviews the 2 See Simpson, A. J., A. W. Kendall, Jr., and J. W. concerning these subgenera of northeast 23 subgenera that have been erected over Orr. 2000. Translations of the original descrip­ Pacific rockfishes and the importance of the years. This review of research, which tions of Sebastes by Georges Cuvier. U.S. Dep. the characters used to distinguish them. includes morphological and genetic studies, Commer., NOAA, Natl. Mar. Fish. Serv., Alaska provides a framework against which to eval­ Fish. Sci. Cent., 7600 Sand Point Way NE, Seat­ Reading the papers of Ayres (1854a, b, uate studies using new genetic techniques. tle WA 98115. Proc. Rep. 2000-09, 24 p. 1855, 1859, 1863a, b, c, d), Gill (1861,

62(2), 2000 1 Figure 1.—Chronology of generic and subgeneric names for Sebastes. Based on Ishida (1984). Generic names are in bold. References below the names propose or support all the names on the line above them.

2 Marine Fisheries Review 1862a, b, 1864), Jordan and Evermann (1896, 1898), Eigenmann and Beeson (1893, 1894), and Cramer (1895) on the subject indicates that more heat than light was generated. For example, by 1862 Gill had erected two genera (Se­ bastodes for Sebastes paucispinis, and Sebastichthys for all other species) for northeast Pacific rockfishes. He based these subgenera on a few morphological characters. Ayres (1863a, d) rejected the characters of Gill, but placed the 11 spe­ cies known to him in two genera: one with smooth heads, for which he used the name Sebastodes, and the other with spiny heads, which he suggested be in­ cluded with the North Atlantic species in Sebastes. In Gill’s 1864 paper “Criti­ cal remarks on the genera Sebastes and Figure 2.—Cumulative number of described and currently recognized species of Sebastodes of Ayres” he stated: “The Sebastes. value of the characters used to distin­ guish the genera Sebastes, Sebastich­ stead of limiting himself to synoptical al fiefdom, the fishes of the North Pa­ thys and Sebastodes are now indeed so tables with minute sub-generic subdi­ cific” (Leviton and Aldrich, 1997, foot­ generally conceded by scientific men, visions. As regards his frequent critical note 16.3, p. 196). that it is unnecessary to further argue in remarks on synonyms, it would be very In 1880, Jordan and Gilbert (1880) their favor. I shall only remark that the useful if he would state whether he ar­ described 7 new species from Califor­ combinations and distinction of forms rived at his conclusions from an exam­ nia, and after trying to place them in by Dr. Ayres are alike unnatural and vi­ ination of typical specimens, but it is the four genera then recognized by Gill, olate all natural affinities . . .” In dis­ not even evident whether he has known they left paucispinis in Sebastodes and cussing Ayres’ ideas on affinities of flat­ the species from autopsy.” Ayres (Fig. stated that: “the groups Sebastosomus fishes, Gill (1864) stated: “Dr. Ayres . 3) was not the only naturalist from San and Sebastomus cannot be maintained . . ideas of affinity are extremely crude Francisco to receive severe criticism as genera distinct from Sebastichthys, and unreliable, nothing can be learned from Gill (Fig. 4): “Of course, Gill had and that, in order to recognize them as from them.” However, in discussing an agenda for this criticism for he ob­ subgenera even, a different distribution this paper Günther (1865) commented: jected to the intrusion of these Califor­ of species must be adopted.” “Dr. Gill would advance ichthyology by nian upstarts, amateurs in his opinion, Eigenmann and Beeson (1893, 1894) giving us serviceable descriptions, in- in what he had carved out as his person- tried to bring some order to the argument

Figure 3.—Photograph of W. O. Ayres, pregnant with interest for him” (Hawkes, reproduced through courtesy of California 1887). Ayres migrated to San Francisco Academy of Sciences Archives. William in 1854 and became a leading member Orville Ayres (1805–1887) was born in and officer in the California Academy of Connecticut and educated at Yale Univer­ Sciences, which met every, or every-other sity to be a medical doctor (Hawkes, 1887). Monday evening in the 1850–1870 period, An early interest in natural history caught to hear of new natural history discoveries the attention, and fostered the friendship of of the area. Besides maintaining a highly­ J. J. Audubon, who named a woodpecker regarded medical practice, Ayres often pre­ (Picus ayresii) in Ayres’ honor stating “I sented descriptions of fishes he came across have named this handsome bird after my at the fish markets, and even presented young and learned friend W. O. Ayres, Esq. information on birds at some of the meet­ who is well known to science as an excellent ings. However “. . . Gill’s ruthless criticism ichthyologist; and who is well conversant of Ayres, finally caused Ayres to give up with the birds of our country” (as seen in research in ichthyology in 1864. Ayres left 1967 republication of Audubon, 1840–1844 San Francisco in 1871 and returned East, [the bird is now considered a hybrid of first to Chicago, Ill., where he suffered seri­ the yellow- and red-shafted flicker]). In ous financial reversals and then in 1878 to 1844, coincident with Audubon’s death, New Haven, Conn., where he re-established convinced that “no field had been ung­ his medical practice and taught at Yale’s leaned in ornithology, [Ayres] turned to medical school” (Leviton and Aldrich, 1997: the water, teeming with attractive life and footnote 3.2, p. 43).

62(2), 2000 3 ian Institution Archives. Record Unit 95, During this work he encountered many of Photograph Collection, 1850s-). Theodore the newly described and undescribed spe­ Nicholas Gill (1837–1914) grew up in New cies of Sebastes from the northeast Pacific York City where early visits to the Fulton and the work of W. O. Ayres on them. As Fish Market spurred his interest in ich­ a result of his studies he published several thyology (Dall, 1916). Early on Gill was descriptions of new species and erected sev­ schooled in the classical languages, with his eral new genera to contain Pacific Sebastes. father intending him to study theology, but In these publications he frequently ques­ this, as well as a job in a law firm, did not tioned Ayres’ work, and eventually pub­ suit him, and his inclination in natural his­ lished (Gill, 1864) a criticism of Ayres tory prevailed. He was granted a scholar­ (Ayres, 1863a) who had taken Gill to task ship to study science in Philadelphia, and for his classification of rockfishes. Ayres from there he went to Washington, D.C., took this criticism so hard that he ceased to where he found a mentor and supporter in publish ichthyological papers. Gill remained S. F. Baird at the Smithsonian Institution. in Washington, D.C., for the rest of his life, Baird published one of Gill’s first papers, on working on fishes, mollusks, mammals, and the fishes of New York, when Gill was only birds. Gill’s strength was in higher order 19 years old (Gill, 1856). Shortly thereaf­ classification rather than in species descrip­ ter Gill made his only extensive field trip tions (“In matters of classification [taxon­ in which he studied the marine and fresh­ omy] he was easily first in the world” Jordan water fishes of the Antilles Islands. Upon (1931)). He had a reputation of being very returning to Washington he settled at the helpful to up-coming students of natural Figure 4.—Photograph of T. N. Gill, Smithsonian and, through Baird’s influence, history, but this did not seem to extend to reproduced with permission (Neg. was appointed to work on fishes collected Ayres. #SA602. Theodore N. Gill. Smithson- during the Northwest Boundary Survey. by publishing a key and diagram of re- ther, Cramer (1895), in his detailed study classic “The Fishes of North and Middle lationships for the eight subgenera they of the cranial osteology of these fishes, America.” In the introduction to their recognized (with mystinus as the sole took strong exception with the impor­ species accounts of Sebastes, Jordan and member of Primospina representing the tance of the first dichotomy (union or Evermann (1898) included from Cramer base from which all other species radi- nonunion of the parietals) in the key of (1895) extensive quotations from his text, ate) (Fig. 5). However, a footnote by the Eigenmann and Beeson (1893). Since and his entire key to species, which they editor (R. Edward Earll) to the title of the Cramer’s work was in essence a rebuttal amended slightly by adding a few species 1894 paper which appeared in the Pro- of the work of Eigenmann and Beeson, Cramer did not consider, and from Ei­ ceedings of the U.S. National Museum he included their entire article as an ap­ genmann and Beeson (1893) their key to stated: “The classification adopted by the pendix to his paper. Cramer (1895) did subgenera. Jordan and Evermann (1898) authors of this paper is based on their not explicitly recognize any subgenera commented on Eigenmann and Beeson’s own peculiar interpretation of the im- for the 32 species of Sebastes he exam­ (1894) work saying: “Messrs. Eigen­ portance of certain structural characters. ined, but his key to species based on cra- mann & Beeson have attempted to sub­ The arrangement and nomenclature pro- nial characters exactly groups the spe­ divide this genus into several subgenera posed here will not be, at present at least, cies into the subgenera eventually used on the basis of cranial characters. Mr. followed by the National Museum.” Fur- by Jordan and Evermann (1898) in their Cramer has given in detail...his reasons for rejecting these proposed genera and for reverting to the sequence of species in Jordan & Gilbert’s Synopsis [1883]. The character especially put forward by Eigenmann & Beeson, that of the con­ tact (not union) of the parietals, seems to us of very slight value, even at a specific distinction.” Consolidation: the Early 1900’s The proliferation of subgeneric names for rockfishes of the Northeast Pacific slowed after the turn of the 20th century. One more was proposed by Jordan and Hubbs (1925), two more by Jordan and Figure 5.—Relationships among subgenera of Sebastes proposed by Eigenmann and Evermann (1927), and one by Hubbs Beeson (1893). (1951). Through the first half of the

4 Marine Fisheries Review 20th century, authors variously lumped, es from both the Atlantic and Pacific split, or ignored subgenera completely. waters of North America. Although in their “Checklist of Fishes of North and Middle America,” Jordan Limited Morphological Studies: et al. (1930) placed 66 species of rock­ the 1970’s fishes they recognized in 16 genera Chen (1971) reviewed Sebastomus, (that is, they elevated the subgeneric the most distinctive subgenus of Se­ names to generic status), most Ameri­ bastes. Occurring in the northeast Pa­ can researchers used only two genera: cific and in the Southern Hemisphere, Sebastes for Atlantic species and Se­ Sebastomus contains 16 species, which bastodes for Pacific species, disregard­ are all characterized by having strong ing the subgenera. head spines, the second anal spine Matsubara (1943) (Fig. 6) presented longer than the third, and a series of six a very detailed analysis of the Scor­ large white blotches (one at the tip of paenidae of the Northwest Pacific. He the opercle and five between the dorsal subdivided the family into 14 subfami­ fin and lateral line [not all are present in lies, and in the Sebastinae he included all species]). Chen (1971) gave detailed four genera, one of which was Se­ accounts of the species and construct­ bastes. He grouped the 30 species of ed a diagram of relationships among Sebastes he considered into 10 subgen­ the 13 then-known species based on a era, 5 of which were newly erected. subjective analysis of about 15 char­ Figure 6.—Photograph of K. Mat­ The five previously established subgen­ acters. He considered phenetic similar­ subara, reproduced with permission era he used had representative species ity among the species in constructing of the American Society of Ichthy­ ologists and Herpetologists. Kiyo­ in the Northeast Pacific also. Matsub­ his diagram. The characters he used matsu Matsubara (1907–1968) was a ara (1943) provided a key to the sub­ were color patterns, body configuration prominent ichthyologist and teacher genera he used, and descriptions based (morphometrics and head spines), and of ichthyology in Japan, who cred­ on a number of osteological, meristic, meristics. He did not specify the out­ ited his interest in fishes to Carl Hubbs (Hubbs, 1969). Matsubara and morphometric characters. He also group for his analysis, or what he con­ published several papers on scor­ concluded that there was no validity sidered to be the primitive states of the paenoid fishes of Japan, capped by in grouping the Atlantic species of Se­ characters he used. Without this infor­ his seminal monograph revising the bastes separately from the Pacific spe­ mation and with the informal discus­ Japanese fishes in this suborder and cies (then considered in Sebastodes), sion of relationships he offered, it is dif­ providing significant insight into the systematics of the group world­ and since Sebastes is the older name it ficult to evaluate his conclusions. Thus, wide (Matsubara, 1943). Consider­ takes precedence. even within this relatively well-known ing the world situation at the time, The early-described Pacific species and cohesive species group, consider­ besides the remarkable ichthyolo­ have fewer dorsal spines and vertebrae able additional work is needed. gical endeavor this monograph rep­ resents, it is amazing that it was pub­ than the Atlantic species, characters Hallacher (1974) characterized the lished in perfect English, in Tokyo. thought important enough to justify structure and points of attachment of Although a species of Sebastes from placing the species from the two oceans swimbladder muscles for 82 species of Japan is named S. matsubarae, it in separate genera. However, species Sebastes, and found two major types was not named after this renowned were later found in the northern Pacific and seven patterns within these types. ichthyologist; rather it was described by Hilgendorf in1880, who named it Ocean and Bering Sea that have verte­ In general, he found little correspon­ after Shinnosuke Matsubara, direc­ bral and dorsal spine counts approach­ dence between species sharing the same tor of the Imperial Fisheries Institute ing those of the Atlantic species, calling muscle patterns and their placement in in Tokyo (Jordan and Starks, 1904). into question the rationale for maintain­ the proposed subgenera (including Se­ Besides his work on scorpaenoid fishes, Matsubara also published sev­ ing the generic distinction. This change bastomus). For example, two similar eral more general volumes on sys­ in thinking is reflected in successive appearing members of Sebastodes (S. tematic ichthyology and papers on editions of the American Fisheries So­ brevispinis and S. paucispinis) had major various other fish groups of Japan. ciety’s “List of Common and Scientific differences in structure. However, all but Although he worked at a time when Names of Fishes from the United States one species of Pteropodus shared the limited access to foreign collections and literature forced many Japanese and Canada.” The first two editions same swimbladder muscle pattern. scientists to consider only local (AFS, 1948; Bailey et al., 1960) rec­ During this period several new spe­ material in their work, Matsubara ommended Sebastes for Atlantic spe­ cies were described or reviewed (e.g. framed his work on scorpaenoids in cies and Sebastodes for Pacific species; Barsukov (1970), Chen (1975), Tsuyuki a larger worldwide context. He was the first to suggest that the genera however, later editions (Bailey et al., and Westrheim (1970)), and some stud­ Sebastes, representing Atlantic spe­ 1970; Robbins et al., 1980; 1991) rec­ ies dealt with the component species cies, and Sebastodes, representing ommended using Sebastes for rockfish­ of some subgenera (e.g. the Sebastes Pacific species, be synonymized.

62(2), 2000 5 Sebastes norvegicus from Cuvier (1829): Plate 23, figure 3 (reproduced by permission of ASHER Rare Books, the Netherlands). This illustration accompanied the original description of the genus Sebastes.

Sebastes marinus (probably S. fasciatus); drawn by H. L. Todd and originally published in Goode and Associates (1884). At the time of this publication only one species of Sebastes was recognized in the northwest Atlantic; however, three species are now known to occur there.

6 Marine Fisheries Review Sebastes ciliatus from the original description of the species by Tilesius (1813). This was the first species of Sebastes to be described from the Northeast Pacific Ocean.

Sebastes aleutianus from Evermann, and Goldsborough (1907). This is one of several fine illustrations of species of Sebastes from Alaska in this early publication.

62(2), 2000 7 vulpes “complex”: Chen and Barsukov rett et al. (1966) examined electropho­ were not related to the geographic dis­ (1976), Sebastiscus: Barsukov and Chen retic patterns of hemoglobin in 27 spe­ tribution of the samples. Among 27 spe­ (1978), and Mebarus: Chen (1985)). cies of Sebastes from off California and cies of Sebastes from the Pacific, 10 had Systematic morphological analysis found that each species, including some unique patterns that allowed them to be of the entire genus during this period morphologically similar species pairs identified based on the muscle protein was limited to Barsukov (1981), who (e.g. Sebastes caurinus-S. vexillaris, S. and enzyme systems used in these stud­ published a complex diagram of inter­ eos-S. chlorostictus), had a distinctive ies. Several species demonstrated dif­ specific relationships of the Sebastinae pattern. They also found shared pat­ ferences in only one enzyme system based on a dubious speciation hypoth­ terns among species in some subgen­ and thus appeared to be closely related esis (Kendall, 1991). Barsukov (1981) era (e.g. Pteropodus, Sebastomus), but to each other (e.g. S. reedi and S. cra­ proposed that there were seven cycles distinct differences among some mem­ meri and S. caurinus, S. aur i culatus and in the evolutionary history of the Sebas­ bers of other subgenera (e.g. Sebas­ S. maliger). There was little variation tinae. “In each one no more than three tosomus). This work was closely fol­ between Atlantic and Pacific Sebastes, descendant species originated sympat­ lowed by a study (Tsuyuki et al., 1968) which were clearly distinct from Sebas­ rically from one ancestral species. Such examining hemoglobin, eye lens, and tolobus and Helicolenus. a triad is characteristic of speciation muscle protein electrophoretic patterns Wishard et al. (1980) used protein with incomplete (mainly biotopic) iso­ in numerous species of Sebastes from electrophoresis to examine within-spe­ lation. A species, once established at both the Atlantic and Pacific, as well as cies relationships among five species intermediate depths, emerges into the representatives of other scorpaeniform of Sebastes from the northeast Pacific. extreme position of another dimension, genera. As in Barrett et al. (1966), spe­ Based on allele frequencies at 21 loci, i.e., it develops ecological and morpho­ cies-specific patterns were found in he­ they found evidence of three popula­ logical features unique for the trio, and moglobins, and they interpreted with­ tions of S. alutus, two of S. pinniger, usually associated with adaptations to in-species variation as evidence of ad­ and one each of S. flavidus, S. goodei, life in open water” (Barsukov, 1981:1). ditional undescribed species. Muscle and S. paucispinis. Based on this premise he reviewed the proteins were not all species-specific, At about the same time, genetic meth­ sequence of geological events that led but they separated the 27 species of Pa­ ods were being used in studies on the to the divergence found in some lineag­ cific Sebastes they examined into four morphologically similar North Atlantic es of these fishes, and related the mor­ subgroups. Comparing these subgroups species of Sebastes (Nefyodov, 1971; phology of living forms to their history, with the subgenera, little correspon­ Naevdal, 1978; Payne and Ni, 1982). ecology, and zoogeography. For exam­ dence is seen. Among the 27 species, A major focus of these studies using ple, in the first cycle he considered the 11 subgenera are represented. Sebastes electrophoresis of hemoglobin and vari­ subfamily Sebastinae to be composed aurora, the sole member of Eosebastes, ous enzyme systems was to confirm the of three genera: the shallow water He­ had a distinct pattern. As in Barrett et presence of more than one sympatric licolenus with 13 species, the interme­ al. (1966), there was some coherence species (S. mentella, S. fasciatus, and diate depth Sebastes with 102 species, among the species of Pteropodus; two S. norvegicus). More recent work on and the deep-water Hozukius with 2 species (S. caurinus and S. nebulosus) Atlantic rockfish has employed genetic species. In the second cycle he felt grouped together as distinct from the techniques to identify specimens, par­ that the genus Sebastes split into three other species. However, another spe­ ticularly juveniles, where more than one subgenera: the shallow water Sebast­ cies of Pteropodus (S. maliger) was species occurs (Nedreaas and Naevdal, iscus with 3 species, the intermediate in a group of 12 species, representing 1991; Rubec et al., 1991), and to exam­ depth Sebastodes with 88 species, and 8 subgenera in which there were no ine population structure within S. nor­ the deep-water Sebastes with 14 spe­ relationships among the species. The vegicus (Nedreaas et al., 1994). cies. More divergence occurred in the muscle and eye lens proteins were dis­ Electrophoretic studies on Pacific Se­ intermediate-depth branch than in the tinct at the generic level. bastes reached a plateau with the work other two during each cycle, and some In the early 1970’s, Johnson et al. of Seeb (1986). She analyzed electro­ branching was not successful (incip­ (1970a, b, 1971, 1972) and Johnson phoretic variations of 28 enzymes in ient species became extinct). Using (1972) conducted a series of investiga­ 48 species of Sebastes and was able to these questionable evolutionary princi­ tions on intraspecific variation in Se­ identify all but a few closely related ples Barsukov (1981) diagramed the re­ bastes alutus and other species of Se­ species pairs on the basis of this anal­ lationships of all known species of the bastes, interspecific variation among Se­ ysis. She produced phenograms based Sebastinae. bastes, and intergeneric variation within on genetic distances among the species. the Scorpaenidae, using electrophoresis As in previous studies (see above), spe­ Genetics Enters the Picture: of muscle proteins and several enzymat­ cies in some subgenera grouped closely the Late 1900’s ic systems. Polymorphisms were found together in these diagrams, while some Starting in the 1960’s, newly devel­ in 11 of the 31 species studied. Three of the branches contained members of oped genetic techniques were applied patterns were seen in S. alutus using two several subgenera. For example, all the to systematic studies of rockfishes. Bar­ enzyme systems, however these patterns members of Zalopyr, Sebastes, and Se­

8 Marine Fisheries Review bastomus that she considered grouped mining the sequence of nucleotides in species of Sebastes had been described together, as did most species of Ptero­ the cytochrome b gene and part of the from the Southern Hemisphere, all ap­ podus, and Sebastodes. However, spe­ control region of the mtDNA molecule parently in Sebastomus. After reviewing cies in Acutomentum, Allosebastes, and (a total of 1,633 bases) in Pacific Se­ the literature (e.g. De Buen, 1960), and Sebastosomus were scattered among bastes, particularly with members of specimens from the west coast of South several lineages within her diagrams. the subgenus Sebastomus. These data America and from South Africa, Chen Further work with allozymes of Pacific were then used to address a number (1971) concluded that more work was rockfish investigated population struc­ of taxonomic and systematic questions. needed, but that he would consider all ture within S. alutus (Seeb and Gunder­ An examination of 14 species of Sebas­ nominal species to be forms of S. cap­ son, 1988: little structure was found), tomus, and 40 other species of Sebastes, ensis, as have others (Norman, 1937; and the applicability of allozymes in showed that the subgenus was mono­ Kong Urbina, 1985). identifying larval and juvenile speci­ phyletic. Low levels of divergence in However, Eschmeyer and Hureau mens (Seeb and Kendall, 1991: allo­ the genetic data indicated recent rapid (1971) considered the Sebastes off the zyme activity and resolution decreased radiation of the subgenus within the last west coast of South America (S. ocula­ with decreasing size of the fish, but was million years (Fig. 7). Among Sebasto­ tus) different from those around south adequate to aid in identification of ju­ mus, S. rosaceus appeared to represent central Atlantic islands and off South veniles and larvae). the oldest lineage, and the rest of the Africa (S. capensis), and suggested that In the 1990’s, newer, more power­ species belonged to one of two clades: more than one species occurs off the ful genetic techniques were applied a northern clade and a southern clade. Pacific coast of South America. Chen to rockfish taxonomic studies in both Generally, the species pairs produced (1971) and Eschmeyer and Hureau the Atlantic and northeast Pacific. by the molecular data were the same (1971) both pointed to the limited dif­ McGauley and Mulligan (1995) ampli­ ones that Chen (1971) found based on ferentiation of Sebastes below the equa­ fied the mitochondrial rRNA (mtDNA) morphological data. tor as evidence of the difficulty for this genes of Sebastes flavidus using poly­ Rocha-Olivares (1998b) used mul­ Northern Hemisphere, temperate and merase chain reaction (PCR) and then tiplex haplotype-specific PCR primers subarctic genus to cross the tropics. used restriction fragment length poly­ to create subgenus and species-specific Based on genetic sequence data, Ro­ morphism (RFLP) analysis to exam­ assays for portions of the cytochrome-b cha-Olivares et al. (1999b) found that ine population structure. They found gene in four species in the subgenus Se­ within the southern clade of Sebas­ essentially no variation in haplotype bastomus. He then screened 250 adults tomus, S. constellatus represented the frequencies among fish collected from representing 31 species of Sebastes that stock that gave rise to the Southern Vancouver Island (British Columbia), had been identified morphologically, Hemisphere species: S. oculatus off Westport (Washington), and Cordell and found that with few exceptions, South America and S. capensis off Bank (California), indicating that gene specimens were correctly assigned to Africa. Further genetic studies (Ro­ flow may be unrestricted within this species using the primers. Larvae were cha-Olivares et al., 1999c) on South­ species throughout its range. They at­ correctly assigned to the subgenus Se­ ern Hemisphere Sebastes demonstrat­ tributed this to long-range dispersal of bastomus, although none was identified ed that the tropical barrier was crossed larvae. However, analysis of a more as a species from which the primers successfully only once, and that three variable region of the mtDNA may were developed. Rocha-Olivares and lineages (cryptic species?) now exist. have shown differences that their analy­ Vetter (1999) also used sequence data to The ancestral species is represented by sis failed to find. Seeb (1998) inves­ examine intraspecific variation within S. capensis which occurs off Tristan da tigated gene flow among S. auricula­ Sebastes helvomaculatus collected at Cuhna and South Africa; this lineage is tus3, S. caurinus, and S. maliger using five sites from Fairweather Bank in the also present in the southwest Atlantic. both allozymes and restriction analy­ Gulf of Alaska to Cordell Bank off There are two lineages of S. oculatus sis of mtDNA and found evidence of California. They found a high degree off the coasts of South America: one introgression between all three species of population genetic diversity which off the Pacific coast and the other off in fish from Puget Sound, indicating could be related to their life history and the Atlantic coast. some level of hybridization. zoogeography. Recent work on the Atlantic species A major step forward in genetic stud­ The application of genetic techniques of Sebastes has used advanced genetic ies occurred as Rocha-Olivares with is well demonstrated by their resolution techniques to investigate relationships others (Rocha-Olivares, 1998a; Rocha- of taxonomic problems associated with among the three morphologically simi­ Olivares et al., 1999a, b) began deter­ Sebastes from the Southern Hemisphere. lar sympatric species (e.g. Bentzen et Sebastes of the Southern Hemisphere al., 1998; Roques et al., 1999). Bentzen 3 Sebastes auriculatus is in the subgenus Auc­ has proven to be an exceptional chal­ et al. (1998) found an unusual number tospina with S. dalli. However, this study and lenge ever since S. capensis was placed of tandemly repeated copies of a sec­ others (e.g. Johnson et al., 1972) indicated it is in the genus by Cuvier and Valenciennes tion of the mtDNA molecule in all three closely related to other shallow-water deep-bod­ ied spiny rockfishes (e.g. S. caurinus) that are in (1829). By the time of Chen’s review of species. The number of tandem repeats Pteropodus. Sebastomus (Chen, 1971), 11 nominal (9–17, mean 12.43) was similar in all

62(2), 2000 9 Figure 7.—Relationships among the species in the subgenus Sebastomus based on genetic data (from Rocha-Olivares et al., 1999b). Time scales represent different assumed rates of evolution. three species, suggesting that the dupli­ individuals to the correct population cichlids and icefishes) in which “spe­ cation might have preceded the diver­ within species. They also concluded, cies flocks” occur. They found that gence of the species. They speculated on the basis of shared alleles, that S. many of the speciation events in Se­ that the Pacific sister species of the At­ norvegicus represents the lineage from bastes occurred in clusters in time, lantic species might have homologous which the other Atlantic species arose, and that these events were far more repeats. They investigated this hypoth­ and that S. fasciatus and S. viviparus ancient than similar events in African esis with S. aleutianus, which is anoth­ are more closely related to each other cichlids. They concluded that rockfish­ er member of the subgenus Sebastes than to the other Atlantic species. es are an ancient species flock, with according to Barsukov (1981), but did As new genetic techniques are de­ significant radiation occurring about 5 not find the tandem repeats. They sug­ veloped, they continue to be applied million years ago (3.6–18 million years gested that other Pacific species could to systematic and taxonomic studies of ago depending on assumptions con­ be examined to determine if they had rockfishes at a variety of levels. For cerning rates of evolution). the tandem repeats and thus might example, Johns and Avise (1998) se­ With the rapid advances in genetic represent the lineage from which the quenced the mitochondrial cytochrome­ techniques that are now occurring, it is Atlantic species arose. Roques et al. b gene from 28 species of rockfish and difficult to predict the limits of what (1999) found that microsatellite data used these data, as well as the allozyme will be known of rockfish taxonomy from eight loci could be used to assign data of Seeb (1986), to construct mo­ and systematics just a few years from individual fish to the correct species lecular phylogenies. They then com­ now. However, as more genomic data more precisely than with other genetic pared these phylogenies with those of are accumulated from these phyloge­ techniques. It was less likely to assign other groups of fishes (Lake Victoria netically interesting fishes, further ad­

10 Marine Fisheries Review Table 1.—Sebastiscus, and Hozukius, and Sebastes subgenera: type species, ranges, numbers of species, and authors.

Subgenus Type species Range No. of species Authors Reference

Acutomentum ovalis NE-NW Pac. 11 Eigenmann and Beeson, 1893 Am. Nat. 27:669 Allosebastes sinensis NE Pac.-Gulf Calif. 14 Hubbs, 1951 Proc. Biol. Soc. Wash. 64:129-130 Auctospina auriculatus NE Pac. 2 Eigenmann and Beeson, 1893 Am. Nat. 27:670 Emmelas glaucus NW Pac. 1 Jordan and Evermann, 1898 Bull. U.S. Natl. Mus. 47:1773 Eosebastes aurora NE Pac. 3 Jordan and Evermann, 1896 Rep. U.S. Comm. Fish Fish. 1895, pt. 21:430 Hatumeus owstoni NW Pac. 1 Matsubara, 1943 Trans. Sigenkagaku Kenkyusyo Hispaniscus rubrivinctus NE Pac. 3 Jordan and Evermann, 1896 Rep. U.S. Comm. Fish Fish. 1895, pt. 21:431 Hozukius emblemarius1 NW Pac. 2 Matsubara, 1934 J. Imp. Fish. Inst. Tokyo 30:199-210. Mebarus inermis NE-NW Pac. 7 Matsubara, 1943 Trans. Sigenkagaku Kenkyusyo:194 Murasoius pachycephalus NW Pac. 1 Matsubara, 1943 Trans. Sigenkagaku Kenkyusyo:235 Neohispaniscus schlegeli NW Pac. 2 Matsubara, 1943 Trans. Sigenkagaku Kenkyusyo:226 Primospina mystinus NE Pac. 1 Eigenmann and Beeson, 1893 Am. Nat. 27:669 Pteropodus maliger NE-NW Pac. 10 Eigenmann and Beeson, 1893 Am. Nat. 27:670 Rosicola pinniger NE Pac. 3 Jordan and Evermann, 1896 Rep. U.S. Comm. Fish Fish. 1895, pt. 21:429 Sebastes norvegicus N Atl. 4 Ascanius, 1772 Claude Philibert, pt. 2 Sebastichthys nigrocinctus NE Pac. 1 Gill, 1862 Proc. Acad. Nat. Sci. Phila. 14:278, 329 Sebastiscus marmoratus NW Pac. 2 Jordan and Starks, 1904 Proc. U.S. Nat. Mus. 27:91-175 Sebastocarus serriceps NE Pac. 1 Jordan and Evermann, 1927 Proc. Calif. Acad. Sci., Ser. 4, 16:507 Sebastocles hubbsi NW Pac. 1 Jordan and Hubbs, 1925 Mem. Carnegie Mus. 10:260. Sebastodes paucispinis NE-NW Pac. 5 Gill, 1861 Proc. Acad. Nat. Sci. Phila. 13:165 Sebastomus rosaceus NE Pac.-S Hemis. 16 Gill, 1864 Proc. Acad. Nat. Sci. Phila. 16:147 Sebastopyr rubberimus NE Pac. 1 Jordan and Evermann, 1927 Proc. Calif. Acad. Sci., Ser. 4, 16:506 Sebastosomus melanops NE Pac. 5 Gill, 1864 Proc. Acad. Nat. Sci. Phila. 16:147 Takenokius oblongus NW Pac. 1 Matsubara, 1943 Trans. Sigenkagaku Kenkyusyo:233 Zalopyr aleutianus NE-NW Pac. 3 Jordan and Evermann, 1898 Bull. U.S. Natl. Mus. 47:1795

1 The species name emblemarius has been misspelled embremarius in several publications including Matsubara (1934, 1943). vances in our understanding of rela­ species in the Southern Hemisphere are rized his views on the relationships tionships among them and their evolu­ in Sebastomus, which also occurs in the within Sebastes, saying “. . . Sebastes tion are assured. northeast Pacific. Some of the subgenera and Sebastiscus of Matsubara (1943) have been proposed as genera, and some are adopted provisionally and subgen­ Present Views groups of subgenera have been con­ era are not used until world-wide review on the Subgenera sidered as genera. No rigorous analy­ of these fishes is carried out.” Although the subgeneric names are sis of the validity of the subgenera has mentioned occasionally in recent litera­ yet been attempted, except for Sebasto­ Challenges ture, all but Sebastomus still lack rigor­ mus (Rocha-Olivares, 1999a). ous definitions. There is also a lack of Several field guides and keys for Where Does Sebastes Belong? agreement concerning the appropriate northeast Pacific Sebastes have been Sebastes is very provisionally placed subgenus for some species (Appendix published (Phillips, 1957; Hitz, 1965; in the order Scorpaeniformes, subor­ II). Some species have been assigned Miller and Lea, 1972; Kramer and der Scorpaenoidei, family Sebastidae, to several different subgenera by vari­ O’Connell, 1988; Orr et al., 2000) subfamily Sebastinae (Nelson, 1994). ous authors through the years. For ex­ which assist greatly in identifying the Much work is yet to be done on the sys­ ample, Allosebastes was erected for Se­ sometimes bewildering array of rock­ tematics of Sebastes, beginning at the bastes sinensis alone by Hubbs (1951); fishes caught even at a single location ordinal level (is the Scorpaeniformes however, Chen (1986) placed 11 other (Table 2). These guides generally rely monophyletic? How does it relate to species in the subgenus, without giving on head spine, meristic, morphomet­ the Perciformes?). The placement of a reason for his action. ric, and color characters that are prob­ Sebastes within the Scorpaeniformes Based on the literature, subgeneric ably not indicative of systematic rela­ is presently uncertain. For example, if names have been assigned to 96 of tionships among species. The subgen­ the Perciformes and Scorpaeniformes the currently recognized species of Se­ era are usually not mentioned in field had a common ancestor, could it have bastes (Table 1). There are 23 subgen­ guides, although Sebastomus is occa­ been similar to Sebastes? Sebastes and era containing from 1 to 16 species. sionally noted since the species are basal perciforms share many character­ Nine subgenera contain only one spe­ quite distinctive externally. istics. The hallmark of the scorpaeni­ cies. Five subgenera have species in Ishida (1984) reviewed what he con­ forms, the suborbital stay is much re­ both the northwest and northeast Pa­ sidered to be the Japanese Sebastinae duced in Sebastes, and, except for the cific Ocean, six have species only in (Sebastes, 26 species; Helicolenus, 3 suborbital stay, Sebastes is more similar the northwest Pacific, and ten are ex­ species; Hozukius, 2 species; and Sebas­ to basal perciforms than are other scor­ clusively in the northeast Pacific. The tiscus, 2 species). He included a careful paeniforms. In other ways, Sebastes is North Atlantic contains a subgenus, Se­ evaluation of all species in these genera the least specialized of the scorpaeni­ bastes, that occurs only there, while the that occur around Japan. He summa­ forms: e.g. head spination is minimal,

62(2), 2000 11 Table 2.—Major field guides to identify species of Northeast Pacific Sebastes.

Reference Geographic area No. of species Presentation

Clemens and Wilby (1946) British Columbia. 20 Extended individual species accounts including airbrush monochrome drawings. Phillips (1957) California, but ranges of some species extend to 49 Key to the species and individual species accounts with monochrome Baja California and Alaska. photographs. Clemens and Wilby (1961) British Columbia. 23 Revision of Clemens and Wilby (1946). Hitz (1965)1 Baja California to the Bering Sea. 53 Species pages arranged by body color and relative length of 2nd and 3rd anal spines. Brief species accounts with black-and-white line sketches and head spine diagrams. Miller and Lea (1972) California, but ranges of some species extend to 58 Brief individual species accounts and a key illustrated with Baja California and Alaska. monochrome sketches showing important characters. Hart (1973) British Columbia. 33 Key to the species and extended individual species accounts with monochrome drawings. Eschmeyer et al. (1983) Coastal waters from the Gulf of Alaska to 58 Individual species accounts and color plates annotated with important Baja California. characters. Kramer and O’Connell (1988) Species that occur off Alaska, includes most species 35 Brief individual species accounts including head spine diagrams, from British Columbia, Washington, and Oregon also. sketches of important characters, and color photographs. Orr et al. (2000) Northeast Pacific Ocean north of Mexico, 66 Format follows Hitz (1965). Includes individual species accounts and including the eastern Bering Sea. color photographs of most species.

1 Hitz (1965) was updated, expanded and reprinted three times in informal editions: in 1977, 1981, and 1991.

Table 3.—Characters used to evaluate genera of Sebastidae, following Ishida, 1994.

Characters States and coding

Number Description Primitive (0) Intermediate (1) Derived (2) Distribution of derived states outside Sebastidae

3 A1 fibers long short Congiopodidae 17 insertion of transversus branched unbranched Congiopodidae and occasional elsewhere dorsalis anterior 41 swimbladder present absent Widespread, especially Scorpaenidae 56 infraorbitals 4 & 5 both present one present both absent Very widespread 64 pharyngobranchials 2 & 3, all separate pharyngobranchials fused all fused Intermediate rare, derived Aploactinidae and Pataecidae upper pharyngeal 70 number of vertebrae 24 more than 24 Widespread except Setarchidae and Scorpaenidae 80 number of hypurals 2+3 1+3 1+2 Derived states widespread 83 scapula and upper radial autogenous fused Widespread 88 supraneural 1 present absent Very widespread

squamation is normal. Is this due to examine the Scorpaenoidei (Tables 3, Table 4.—Characters and character states for analysis convergence or a common ancestor? 4). All members of the family have de­ of Sebastidae, following Ishida, 1994. Basic unresolved questions regarding rived states of two characters relative Characters1 the origin and relationships of Sebastes to their states in lower percoids, the Genus 3 17 41 56 64 70 80 83 88 and its close relatives include: was their presumed outgroup of scorpaenoids: Sebastes 1 0 0 0 0 1 2 1 0 ancestor tropical or boreal, and was the shortening of a1 fibers with the Sebastiscus 1 0 0 0 0 1 2 1 0 it demersal or pelagic? Does Sebastes lengthening of the associated maxillary Hozukius 1 0 0 0 0 1 2 1 0 Helicolenus 1 0 1 0 0 1 2 1 0 represent the point of divergence that tendon (character 3), and an increase in Adelosebastes 1 0 1 0 0 1 2 1 0 led the demersal-boreal scorpaeniforms the number of vertebrae from 24 (char­ Sebastolobus 1 0 1 0 0 1 2 0 0 Trachyscorpia 1 0 1 1 0 1 0 0 0 away from the pelagic-tropical perci­ acter 70) (Table 3). Plectrogenium 1 1 1 1 1 1 0 0 1 forms, or does the genus represent an Within the Sebastidae, the genera end point in scorpaeniform evolution Sebastes, Sebastiscus, and Hozukius 1 0=primitive, 1=intermediate, 2=derived. (Washington et al., 1984a)? Although share identical character states, result­ these questions may eventually be re­ ing in an unresolved trichotomy, which reversal (in that the rest of the family solved using cladistic methodology, the can be given the subfamilial name Se­ lacks them, which is the presumed de­ diversity and numbers of species in bastinae. This subfamily has the de­ rived state). If Hozukius is found to be both the perciforms and scorpaeniforms rived states of four characters and viviparous like Sebastes and Sebastis­ may continue to deter their resolution. shares the derived states of two of these cus, then viviparity would be another Within the suborder Scorpaenoidei, characters with all of the other genera derived character to support this sub­ Ishida (1994) considered the Sebastidae in Sebastidae. However, they do not family. If Hozukius is found to be ovip­ to contain eight genera. Some members possess uniquely derived states of any arous, it may be considered the prim­ of the Sebastidae have derived states of characters. They are united in possess­ itive sister genus of Sebastes and Se­ 9 of the 95 characters that he used to ing swimbladders, which is a character bastiscus within the subfamily.

12 Marine Fisheries Review Table 5.—Ecological correlates of characters of Sebastes. these characters and Emmelas, which larger and slightly separate from the Demersal Pelagic lives in the northern part of the range, nuchal spine (Moser et al., 1977; Moser, Character state state has higher counts. The species of Se­ 1996). There is a tendency for early de­ Head spines strength Strong Weak bastomus, which live primarily in the velopment of pectoral fins in scorpaeni­ Head spine numbers Many Few Jaw length Lower > upper Equal southern part of the range of Sebastes forms, and in some groups the pectoral Interorbital shape Concave Convex in the northeast Pacific, including the fins are quite pronounced in the larvae. Lower pectoral rays Thick Not thick Gill raker length Short Long Gulf of California, and the Southern Pronounced pectoral fins are seen in Se­ Jaws scaled Yes No Base of skull Curved Flat Hemisphere, have lower medial fin ray bastolobus, Scorpaenodes, Ectrepose­ Mesethmoid process Strong Weak counts than most other Sebastes. The bastes and some species of Sebastes Color Red Black Atlantic species of Sebastes have the (e.g. S. levis). Larvae of Helicolenus de­ Low latitude High latitude highest vertebral counts in the genus, velop a spongy mass of tissue anterior­ state state and presumably they originated from ly along the dorsal midline. Once the Vertebrae <29 >28 species that traversed the Arctic Ocean larvae of more scorpaenoids become Dorsal spines <14 14–15 Dorsal rays <14 15–16 from the Pacific. known, such characters will prove in­ Anal rays 5 9 valuable in systematic studies of the Principal caudal rays 13 15 Can Early Life History Pectoral rays 17–18 >18 group. Information Be Applied Within the genus Sebastes, the pre­ to the Problem? flexion-extrusion larvae of most spe­ What Morphological Characters Scorpaenoids display a variety of re­ cies are known (Matarese et al., 1989; Can Be Used for productive patterns, but the use of in­ Moser, 1996). These can be obtained Cladistic Analysis of Sebastes? formation on reproduction in system­ fairly easily, and their identity known, To be used in a cladistic analysis, atic studies is limited, since the pat­ by collecting pregnant females and ex­ morphological characters must be in­ terns are unknown in several genera truding the larvae. Aside from some dependent of each other and nonadap­ (Washington et al., 1984b). While most shared characteristics among preflex­ tive. However, many of the characters scorpaenoids share ovoviviparity with ion larvae of species of Sebastomus, used to identify Sebastes and used in most other teleosts, Sebastes, Sebast­ the appearance of these larvae seems early attempts to establish relationships iscus, and possibly Helicolenus share to offer little taxonomic or systematic among the species are correlated with the unique pattern of internal fertil­ promise. There is considerable overlap each other and are adaptive for either ization, intraovarian embryological de­ in characters among species in these pelagic or demersal existence (Table 5). velopment with maternal contribution larvae and considerable within-species There is a basic dichotomy in Se­ to nutrition, and production of many variation. bastes ecology: species either live in (~105) newly hatched, undeveloped Later-stage larvae of Sebastes are the water column or on the bottom. larvae. The reproductive pattern of Ho­ distinctive from larvae of other fishes, This is not an absolute; there is a zukius is unknown, although the anal but many share characters with each cline between these two basic condi­ anatomy is that of a live-bearer (Ida4). other that prevent identifying them to tions. Bottom-dwelling species have As opposed to the widespread pattern of species in all but a few cases. Larval the lower jaw protruding, eyes close producing individual planktonic eggs, characters that may be useful in tax­ together with a concave interorbital several scorpaenid genera (Sebastolo­ onomy and systematics include body area, heavy armature, thickened pecto­ bus, Pterois, Scorpaena) extrude eggs shape, pigment patterns, and the de­ ral rays, short gill rakers, scaled jaws, in a floating bilobed gelatinous egg velopment and subsequent reduction or and a curved skull base, while pelagic mass. A single oil globule is present in loss of head spines (Kendall, 1991). species have opposite states for all of eggs of Pterois and Sebastolobus, while As with early larvae, the later larvae these characters. Convergence toward none is present in the eggs of Scorpae­ of Sebastomus share some characters these character states is expected in na or Dendrochirus. that distinguish them from other Se­ species occupying these niches regard­ The larvae of many scorpaeniforms, bastes larvae, but within the subgenus, less of their ancestry. including the scorpaenoids, have pari­ the larvae of many species are quite Another factor to consider when in­ etal and other head spines, which are similar. It appears that larvae of Ptero­ terpreting characters is that meristic not seen in larvae of other fishes. In Se­ podus may also share some characters, characters tend to show a latitudinal bastolobus and Scorpaenodes the pari­ but the larvae of several species are still cline: the closer the species lives to etal ridge is enlarged and ends in a bifid unknown. A few other species have dis­ the pole, the higher the value (Jordan’s spine, with the posterior part (the nuchal tinctive larvae (e.g. Sebastes paucispi­ Rule: see Lindsey, 1988). This basic spine) being predominant, whereas in nis, S. jordani, S. melanostomus, and S. pattern is seen in medial fin ray and ver­ Sebastes, Sebastiscus (Okiyama, 1988), levis), but their morphology does not tebral counts among species of rock­ and Helicolenus the parietal spine is seem to be related to their subgeneric fishes (Table 5). For example, Sebas­ placement. There are some intriguing tiscus, which lives farther south than 4 Ida, Hitoshi, Kitasato Univ., Sanriku, Jpn., per­ similarities among the larvae of some most other taxa, has low counts for all sonal commun., 20 January 1999. species of Sebastes that may indicate

62(2), 2000 13 relationships not otherwise suggested able through genetics, need to be evalu­ Lindsey, C. R. Robbins, and W. B. Scott. 1970. A list of common and scientific names (e.g. the larvae of S. jordani, S. alutus, ated more rigorously. With the incredi­ of fishes from the United States and Canada, and S. polyspinis from the Pacific all ble size of the genus and its wide distri­ third ed. Am. Fish. Soc. Spec. Publ. 6, 150 p. look quite similar to the larvae of the butional range, this will be an enormous Barrett, I., J. Joseph, and H. G. Moser. 1966. Elec­ trophoretic analysis of hemoglobins of Cal­ Atlantic species of Sebastes). task. ifornia rockfish (genus Sebastodes). Copeia As juveniles, Sebastes are morpho­ 1966:489–494. logically quite different from the larvae Acknowledgments Barsukov, V. V., and L.-C. Chen. 1978. Review of the subgenus Sebastiscus (Sebastes, Scorpae­ and the adults. Some species reside in I am pleased to express appreciation nidae) with a description of a new species. J. the water column, some associate with to the following people for their help Ichthyol. 18:179–193. ______. 1970. Species composition of the flotsam or school, and some take up with this work. Bruce Collette, NMFS genus Sebastes in the North Pacific. Descrip­ a demersal existence (Moser and Boe­ Systematics Laboratory, supplied draft tion of a new species. Dokl. Acad. Nauk SSSR hlert, 1991). The observed pigment pat­ copies of work in progress on North At­ Ser. Biol. 195:994–997. ______. 1981. A brief review of the subfamily terns of juveniles may be adaptive for lantic rockfishes and rockfish specimens Sebastinae. J. Ichthyol. 18:179–193. these various modes of existence, so from the North Atlantic for study. Abby Bentzen, P., J. M. Wright, L. T. Bryden, et caution is advised when using them in al. 1998. Tandem repeat polymorphism and J. Simpson, formerly of the School heteroplasmy in the mitochondrial control systematic analysis. of Fisheries, University of Washing­ region of red fishes (Sebastes: Scorpaenidae). Clearly the early life history of scor­ ton, Seattle, kindly translated Cuvier’s J. Heredity. 89:1–7. Chen, L.-C. 1971. Systematics, variation, distribu­ paenoids in general and Sebastes in par­ original description of Sebastes from tion, and biology of rockfishes of the subgenus ticular offers a large number of char­ French. Tony Gharrett, University of Sebastomus (Pisces, Scorpaenidae, Sebastes). acters to apply to systematic studies. Alaska; Geoff Moser, SWFSC; and Jay Bull. Scripps Inst. Oceanogr. 18:1–115. ______. 1975. The rockfishes, genus Sebastes However, until the reproductive pat­ Orr, AFSC, lent much needed encour­ (Scorpaenidae), of the Gulf of California, terns and early life history series of agement for this project and reviewed including three new species with a discussion more species are described, this poten­ of their origin. Proc. Calif. Acad. Sci. Ser. 4. an earlier draft of the manuscript. Mike 40:109–141. tial cannot be realized. Canino, AFSC, and Paul Bentzen, Uni­ ______. 1985. A study of the Sebastes iner­ versity of Washington, reviewed an ear­ mis species complex, with delimination of the Conclusions subgenus Mebarus (Pisces, Scorpaenidae). J. lier draft also. Taiwan Mus. 38:23–37. After a period of discovering the in­ ______. 1986. Meristic variation in Sebastes credible diversity of Sebastes in the Literature Cited (Scorpaenidae) with an analysis of character AFS. 1948. A list of common and scientific association and bilateral pattern and their sig­ middle 1800’s, the history of the sys­ names of better known fishes of the United nificance in species separation. U.S. Dep. tematics of the genus was marked by States and Canada. Am. Fish. Soc. Spec. Publ. Commer., NOAA Tech. Rep. NMFS 45, 17 p. bitter debate in the late 1800’s over re­ 1, 58 p. ______and V. V. Barsukov. 1976. A study of Ascanius, P. 1772. Icones rerum naturalium, the western north Pacific Sebastes vulpes spe­ lationships among the species. With few ou figures enluminées d’histoire naturelle du cies complex (Scorpaenidae) with description exceptions (e.g. S. carnatus-S. chrys­ norde. Claude Philibert, Copenhagen, pt. 2. of a new species. Jpn. J. Ichthyol. 23:1–8. omelas-S. atrovirens, and S. ciliatus) Audubon, J. J. 1840–1844. The birds of America. Clemens, W. A., and G. V. Wilby. 1946. Fishes of Republished 1967. Dover Publ., N.Y., 7 vol. the Pacific coast of Canada. Fish. Res. Board the species of Sebastes are now well­ Ayres, W. O. 1854a. [Descriptions of new fishes Can., Bull. 68, 368 p. defined, and few new ones are being de­ from California.] Proc. Calif. Acad. Nat. Sci. ______and ______. 1961. Fishes of the 1:5–7. Pacific coast of Canada. Fish. Res. Board scribed (Kim and Lee, 1994; Eitner et ______. 1854b. [Descriptions of new fishes Can., Bull. 68, 2nd ed., 443 p. al., 1999). However, little recent work from California.] Proc. Calif. Acad. Nat. Sci. Cramer, F. 1895. On the cranial characters of has been done on the genus as a unit, 1:3–32. (Repr. 1873 in 2nd ed. Proc. Calif. the genus Sebastodes (rock-fish). Proc. Calif. Acad. Nat. Sci.) Acad. Sci., ser. 2, 5(1):573–610, pl. 57–70. and its systematics remain in a confused ______. 1855. New species of California fishes. Cuvier, G. 1829. Le Règne animal, distribuè state, with over 100 species, and very Proc. Boston Soc. Nat. Hist. 5:94–103. d’après son organisation, pour servir de little obvious structure within the genus. ______. 1859. [Descriptions of new species of base à l’histoire naturelle des animaux et fishes.] Proc. Calif. Acad. Nat. Sci. 2:25–32. d’introduction à l’anatomie comparèe. Ed. 2, What other fish genus contains so many ______. 1863a. Notes on the sebastoid fishes v. 2:I–xvii + 1–532. species, but with such little indication occurring on the coast of California, U.S.A. ______and A. Valenciennes. 1829. Histoire Proc. Zool. Soc. Lond. 26:390–402. naturelle des poissons. Tome quatrième. Livre of structure within it? Beyond the fairly ______. 1863b. [Note on Sebastodes rosaceus quatrième. Des acanthoptérygiens à joue cohesive group Sebastomus and a few and Sebastes ruber.] Proc. Calif. Acad. Nat. cuirassée. v. 4:i–xxvi + 2 pp. + 1–518, Pls. species pairs (e.g. Sebastes fasciatus/S. Sci. 2:207. 72–99, 97 bis. ______. 1863c. [Descriptions of Sebastodes Dall, W. H. 1916. Theodore Nicholas Gill. Bio­ mentella, S. babcocki/S. rubrivinctus, S. flavidus and Sebastodes ovalis.] Proc. Calif. graphical memoirs. Natl. Acad. Sci. 8:313– serranoides/S. flavidus), practically any Acad. Nat. Sci. 2:209–211. 343. two species look as similar to each other ______. 1863d. [Remarks in relation to the De Buen, F. 1960. Nota preliminar sobre los fishes of California which are included in peces del genero Sebastodes en la fauna Chil­ as to any other species. Cuvier’s genus Sebastes.] Proc. Calif. Acad. ena. Rev. Chil. Hist. Nat. 55:3–26. In order to progress in our under­ Nat. Sci. 2:211–218. Eigenmann, C. H., and C. H. Beeson. 1893. Pre­ Bailey, R. M., E. A. Lachner, C. C. Lindsey, C. liminary note on the relationship of the spe­ standing of the relationships within this R. Robbins, P. M. Roedel, W. B. Scott, and cies usually united under the generic name interesting, and ecologically and eco­ L. P. Woods. 1960. A list of common and sci­ Sebastodes. Am. Nat. 27:668–671. nomically important group of fishes, entific names of fishes from the United States ______and ______. 1894. A revision of and Canada, second ed. Am. Fish. Soc. Spec. the fishes of the subfamily Sebastinae of the traditional gross morphological as well Publ. 2, 102 p. Pacific coast of America. Proc. U.S. Natl. as novel characters, such as those avail­ ______, E. S. Herald, E. A. Lachner, C. C. Mus. 17:375–407.

14 Marine Fisheries Review Eitner, B., C. Kimbrell, and R. Vetter. 1999. glycerophosphate dehydrogenase in Pacific classes, ordines, genera, species, cum charac­ Sebastes moseri (Scorpaeniformes: Scorpae­ ocean perch (Sebastodes alutus). J. Fish. Res. teribus, differentiis, synonymis, locis. Tomus nidae): A new rockfish from the eastern North Board Can. 27:943–945. 1. Editio decima, reformata.) Laurentii Salvii. Pacific. Copiea 1999:85–92. ______, ______, and ______. 1970b. Holmiae, v. 1:1–ii + 1–824. Eschmeyer, W. N., and J. C. Hureau. 1971. Interspecific variation of tetrazoliurn oxidase ______. 1761. Fauna Svecica sistens anima­ Sebastes mouchezi, a senior synonym of in Sebastodes (rockfish). Comp. Biochem. lia Sveciae Regni:... Editio Altera. Laurentii Helicolenus tristanensis, with comments on Physiol. 37:281–285. Salvii, Stockholm. Sebastes capensis and zoo-geographical con­ ______, ______, and ______. 1971. Matarese, A. C., A. W. Kendall, Jr., D. M. Blood, siderations. Copeia 1971:576–579. Phosphoglucomutase polymorphism in Pacific and B. M. Vinter. 1989. Laboratory guide to ______, E. S. Herald, and H. Hammann. 1983. A ocean perch, Sebastes alutus. Comp. Bio­ early life history stages of northeast Pacific field guide to Pacific coast fishes of North Amer­ chem. Physiol. 31B:285–290. fishes. U.S. Dep. Commer., NOAA Tech. Rep. ica. Houghton Mifflin Co., Boston, 336 p. ______, ______, and ______. 1972. NMFS Circ. 80, 656 p. Evermann, B. W., and E. L. Goldsborough. 1907. Electrophoretic investigation of the family Matsubara, K. 1934. Studies on the scorpaenoid The fishes of Alaska. Bull. U.S. Bur. Fish. Scorpaenidae. Fish. Bull. 70:403–413. fishes of Japan. I. Descriptions of one new 26:219–360. Jordan, D. S. 1887. A catalogue of the fishes genus and five new species. J. Imp. Fish. Inst. Fernholm, B., and A. Wheeler. 1983. Linnaean known to inhabit the waters of North Amer­ Tokyo 30:199–210. fish specimens in the Swedish Museum of ica, north of the Tropic of Cancer, with notes ______. 1943. Studies on the scorpaenoid Natural History, Stockholm. Zool. J. Linn. on species discovered in 1883 and 1884. Rep. fishes of Japan. Anatomy, phylogeny and tax­ Soc. 78:199–286. U.S. Comm. Fish Fish.1885 13:789–973. onomy, I, II. Trans. Sigenkagaku Kenkyusyo Gill, T. 1856. On the fishes of New York. Smith­ ______. 1931. Gill, Theodore Nicholas. In No. 1, 2, Tokyo, 486 p. son. Inst. Annu. Rep. 253–269. A. Johnson and D. Malone (Editors), Diction­ McGauley, K., and T. J. Mulligan. 1995. Poly­ ______. 1861. Notes on some genera of fishes ary of American Biography 7, p. 285–286. merase chain reaction restriction fragment of the western coast of North America. Proc. Charles Scribner’s Sons, N.Y. length polymorphism analysis of mitochon­ Acad. Nat. Sci. Phila. 13:164–168. ______and B. W. Evermann. 1896. A check drial rRNA genes from yellowtail rockfish. J. ______. 1862a. Notice of a collection of the list of the fishes and fish-like vertebrates of Fish. Biol. 47:744–747. fishes of California presented to the Smithso­ North and Middle America. Rep. U.S. Comm. Miller, D. J., and R. N. Lea. 1972. Guide to the nian Institution by Mr. Samuel Hubbard. Proc. Fish Fish. 1895, pt. 21:207–584. coastal fishes of California. Calif. Dep. Fish Acad. Nat. Sci. Phila. 14:274–282. ______and ______. 1898. The fishes of Game, Fish Bull. 157, 249 p. ______. 1862b. Notes on some genera of North and Middle America. Bull. U.S. Natl. Moser, H. G. 1996. Scorpaenidae: scorpionfishes fishes of western North America. Proc. Acad. Mus. 47:1241–2183. and rockfishes. In H. G. Moser (Editor), The Nat. Sci. Phila. 14:329–332. ______and ______. 1927. New genera and early stages of fishes in the California Current ______. 1864. Critical remarks on the genera species of North American fishes. Proc. Calif. region, p. 733–795. CalCOFI Atlas 33. Sebastes and Sebastodes of Ayres. Proc. Acad. Acad. Sci., Ser. 4, 16:501–507. ______, E. H. Ahlstrom, and E. M. Sandknop. Nat. Sci. Phila. 16:145–147. ______, ______, and H. W. Clark. 1930. 1977. Guide to the identification of scorpi­ Goode, G. B., and a staff of Associates. 1884. The Check list of the fishes and fish-like verte­ onfish larvae (family Scorpaenidae) in the fisheries and fishery industries of the United brates of North and Middle America north eastern Pacific with comparative notes on spe­ States. Section I, Atlas of Plates, Plate 75. of the northern boundary of Venezuela and cies of Sebastes and Helicolenus from other Günther, A. 1865. Pisces. The record of the zoo­ Colombia. Rep. U.S. Comm. Fish Fish. 1928, oceans. U.S. Dep. Commer., NOAA Tech. logical literature. Zool. Rec. 1864:133–188. 670 p. Rep. NMFS Circ. 402, 71 p. Hallacher, L. E. 1974. The comparative morphol­ ______and C. H. Gilbert. 1880. Description ______and G. W. Boehlert. 1991. Ecology ogy of extrinsic gasbladder musculature in the of seven new species of sebastoid fishes, from of pelagic larvae and juveniles of the genus scorpionfish genus Sebastes (Pisces: Scorpae­ the coast of California. Proc. U.S. Natl. Mus. Sebastes. Environ. Biol. Fish. 30:203–224. nidae). Proc. Calif. Acad. Sci. 40:59–86. 3:287–298. Naevdal, G. 1978. Differentiation between mari­ Hart, J. L. 1973. Pacific fishes of Canada. Fish. ______and ______. 1883. Synopsis of the nus and mentella types of redfish by electro­ Res. Board Can., Bull. 80, 740 p. fishes of North America. Bull. U.S. Natl. Mus. phoresis of haemoglobins. Fiskeridir. Skr. Ser. Hawkes, W. W. 1887. W. O. Ayres, M.D., New 16:1–1018. Havunders. 16:359–308. Haven. Obituaries. Proc. Connecticut Med. ______and C. L. Hubbs. 1925. Record of Nedreaas, K., and G. Naevdal. 1991. Identi­ Soc. 4:174–176. fishes obtained by David Starr Jordan in Japan, fication of 0- and I-group redfish (genus Hilgendorf, F. M. 1880. Uebersicht über die 1922. Mem. Carnegie Mus. 10:93–346. Sebastes) using electrophoresis. ICES J. Mar. Japanischen Sebastes-Arten. Gesellschaft ______and E. C. Starks. 1904. A review of Sci. 48:91–99. Naturforschenden Freunde zu Berlin, Sizungs­ the scorpaenoid fishes of Japan. Proc. U.S. ______, T. Johansen, and G. Naevdal. 1994. berichten 168–172. Natl. Mus. 27:91–175. Genetic studies of redfish (Sebastes spp.) from Hitz, C. R. 1965. Field identification of the north­ Kendall, A. W., Jr. 1991. Systematics and identi­ Icelandic and Greenland waters. ICES J. Mar. eastern Pacific rockfish (Sebastodes). U.S. fication of larvae and juveniles of the genus Sci. 51:461–467. Dep. Inter., Fish Wildl. Serv., Bur. Commer. Sebastes. Environ. Biol. Fish. 30:173–190. Nefyodov, G. N. 1971. Serum haptoglobins in the Fish., Circ. 203, 58 p. Kim, I. S., and W. O. Lee. 1994. A new species marinus and mentella types of North Atlantic Hubbs, C. L. 1951. Allosebastes, new subgenus of the genus Sebastes (Pisces; Scorpaenidae) redfish. Rapp. P-v. Reun. Cons. Int. Explor. for Sebastodes sinensis, scorpaenid fish of the from the Yellow Sea, Korea. Korean J. Zool. Mer 161:126–129. Gulf of California. Proc. Biol. Soc. Wash. 37:409–415. Nelson, J. S. 1994. Fishes of the world, third edi­ 64:129–130. Kong Urbina, I. 1985. Revision de las especies tion. John Wiley & Sons, N.Y., 624 p. ______. 1969. Kiyomatsu Matsubara chilenas de Sebastes (Osteichthyes, Scor­ Norman, J. R. 1937. Coast fishes. Part II. The Pata­ 1907–1968. Copeia 1969:420–422. paeniformes, Scorpaenidae). Estud. Oceanol. gonian Region. Discovery Rep. 16:1–150. Ishida, M. 1984. Taxonomic study of the sebastine 4:21–75. Okiyama, M. (Editor). 1988. An atlas of the fishes in Japan and its adjacent waters. M.S. Kramer, D. E., and V. M. O’Connell. 1988. Guide early stage fishes in Japan. Tokai Univ. Press, thesis, Univ. Hokkaido, Hakodate, 245 p. to northeast Pacific rockfishes. Univ. Alaska Tokyo, 1154 p. ______. 1994. Phylogeny of the suborder Scor­ Mar. Advis. Bull. 25, 78 p. Orr, J. W., M. A. Brown, and D. C. Baker. 2000. paenoidei (Pisces: Scorpaeniformes). Bull. Leviton, A. E., and M. L. Aldrich. 1997. The Guide to the rockfishes (Scorpaenidae) of the Nansei Natl. Fish. Res. Inst. 27:1–112. California Academy of Sciences: a narrative genera Sebastes, Sebastolobus, and Adelose­ Johns, G. C., and J. C. Avise. 1998. Tests for history: 1853–1906. Calif. Acad. Sci., San bastes of the northeast Pacific Ocean, second ancient species flocks based on molecular Francisco. edition. U.S. Dep. Commer., NOAA Tech. phylogenetic appraisals of Sebastes rock­ Lindsey, C. C. 1988. Factors controlling meris­ Memo. NMFS-AFSC-117, 47 p. fishes and other marine fishes. Evolution tic variation. In W. S. Hoar and D. J. Randall Payne, R. K., and I. H. Ni. 1982. Biochemical 52:1135–1146. (Editors), Fish physiology, vol. 11. The phys­ population genetics of redfishes (Sebastes) off Johnson, A. G. 1972. An electrophoretic inves­ iology of developing fish. Part B. Viviparity Newfoundland. J. Northwest Atl. Fish. Sci. tigation of the family Scorpaenidae. Ph.D. and posthatching juveniles, p. 197–274. Acad. 3:169–172. thesis, Univ. Wash., Seattle, 65 p. Press, San Diego. Phillips, J. B. 1957. A review of the rockfishes of ______, F. M. Utter, and H. O. Hodgins. Linnaeus, C. 1758. Systema naturae, Ed. X. (Sys­ California (family Scorpaenidae). Calif. Dep. 1970a. Electrophoretic variants of L-alpha tema naturae per regna tria naturae, secundum Fish Game, Fish Bull. 104, 158 p.

62(2), 2000 15 Robbins, C. R., R. M. Bailey, C. E. Bond, J. R. zoogeography of the rockfish subgenus Sebas­ Steindachner, F., and L. Döderlein. 1884. Beiträge Brooker, E. A. Lachner, R. N. Lea, and W. B. tomus (Sebastes, Scorpaenidae) based on mito­ zur Kentniss der Fisch Japan, I(IV). Denkshr. Scott. 1980. A list of common and scientific chondrial cytochrome b and control region Akad. Wiss. Wien. 171–212. names of fishes from the United States and sequences. Mol. Phylogenet. Evol. 11:441–458. Tilesius, W. G. von. 1813. Iconum et descrip­ Canada, fourth ed. Am. Fish. Soc. Spec. Publ. ______, ______, and ______. 1999c. tionum piscium Camtschaticorum continuatio 12, 173 p. Cryptic species of rockfishes (Sebastes: Scor­ tertia tentamen monographiae generis Agoni ______, ______, ______, ______, paenidae) in the Southern Hemisphere inferred blochiani sistens. Mem. Acad. Imp. Sci., St. ______, ______, and ______. from mitochondrial lineages. J. Hered. 90: Petersb. (for 1881):406–478. 1991. Common and scientific names of fishes 404–411. Tsuyuki, H., E. Roberts, R. H. Lowes, W. Had­ from the United States and Canada, fifth ed. Roques, S., P. Duchesne, and L. Bernatchez. away, and S. J. Westrheim. 1968. Contribution Am. Fish. Soc. Spec. Publ. 20, 183 p. 1999. Potential of microsatellites for indi­ of protein electrophoresis to rockfish (Scor­ Rocha-Olivares, A. 1998a. Molecular evolution, vidual assignment: the North Atlantic redfish paenidae) systematics. J. Fish. Res. Board systematics, zoogeography and levels of intra­ (genus Sebastes) species complex as a case Can. 25:2477–2501. specific genetic differentiation in the species of study. Molec. Ecol. 8:1703–1717. ______and S. J. Westrheim. 1970. Analysis the antitropical subgenus Sebastomus, Sebastes Rubec, P. J., J. M. McGlade, B. L. Trottier, et of the Sebastes aleutianus-S. melanostomus (Scorpaeniformes, Teleostei) using mitochon­ al. 1991. Evaluation for methods for separa­ complex and description of a new scorpaenid drial DNA sequence data. PhD Dissertation. tion of Gulf of Saint-Lawrence beaked red­ species, Sebastes caenaematicus, in the north­ University of California. San Diego, 240 p. fishes, Sebastes fasciatus and S. mentella: east Pacific Ocean. J. Fish. Res. Board Can. ______. 1998b. Multiplex haplotype-specific malate dehydrogenase mobility patterns com­ 27:2233–2254. PCR: a new approach for species identifica­ pared with extrinsic gasbladder muscle pas­ Washington, B. B., W. N. Eschmeyer, and K. tion of the early life stages of rockfishes of the sages and anal fin ray counts. Can. J. Fish. M. Howe. 1984a. Scorpaeniformes: Relation­ species-rich genus Sebastes Cuvier. J. Exp. Aquat. Sci. 48:640–660. ships. In H. G. Moser, W. J. Richards, D. M. Mar. Biol. Ecol. 231:279–290. Seeb, L. W. 1986. Biochemical systematics and Cohen, M. P. Fahay, A. W. Kendall, Jr., and ______and R. D. Vetter. 1999. Effects of evolution of the scorpaenid genus Sebastes. S. L. Richardson (Editors), Ontogeny and sys­ oceanographic circulation on the gene flow, PhD Dissert., Univ. Wash., Seattle, 176 p. tematics of fishes, p. 438–447. Am. Soc. Ich­ genetic structure, and phylogeography of the ______. 1998. Gene flow and introgression thyol. Herpetol. Spec. Publ. 1. rosethorn rockfish (Sebastes helvomaculatus) within and among three species of rockfishes, Washington, B. B., H. G. Moser, W. A. Laroche, Can. J. Fish. Aquat. Sci. 56:803–813. Sebastes auriculatus, S. caurinus, and S. and W. J. Richards. 1984b. Scorpaeniformes: ______, C. A. Kimbrell, B. J. Eitner, and R. maliger. J. Heredit. 89:393–403. development. In H. G. Moser, W. J. Richards, D. Vetter. 1999a. Evolution of a mitochondrial ______and D. R. Gunderson. 1988. Genetic D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., cytochrome b gene sequence in the species­ variation and population structure of Pacific and S. L. Richardson (Editors), Ontogeny and rich genus Sebastes (Teleostei, Scorpaenidae) ocean perch (Sebastes alutus). Can. J. Fish. systematics of fishes, p. 405–428. Am. Soc. and its utility in testing the monophly of the Aquat. Sci. 45:78–88. Ichthyol. Herpetol. Spec. Publ. 1. subgenus Sebastomus. Mol. Phylogenet. Evol. ______and A.W. Kendall, Jr. 1991. Allozyme Wishard, L. N., F. M. Utter, and D. R. Gunderson. 11:426–440. polymorphisms permit the identification of 1980. Stock separation of five rockfish species ______, R. H. Rosenblatt, and R. D. Vetter. larval and juvenile rockfishes of the genus using naturally occurring biochemical genetic 1999b. Molecular evolution, systematics, and Sebastes. Environ. Biol. Fish. 30:191–201. markers. Mar. Fish. Rev. 42(3–4):64–71.

16 Marine Fisheries Review Appendix I arated and combined under the genus fered; see description of type species in Sebastosomus . . .” Jordan and Evermann (1898). Original Definitions of Sebastes, Acutomentum Eigenmann and Beeson Rosicola Jordan and Evermann (1896), Hozukius, and Sebastiscus, (1893), type species: S. ovalis: Taken type species: S. pinniger: No descrip­ and the Subgenera of Sebastes from their key: “Parietals meeting above tion of the subgenus offered; see de­ Sebastes Cuvier (1829), type species: the supra-occipital. Lower jaw much scription of type species in Jordan and S. norvegicus: “Have all the characters projecting; head broad, the skull usu­ Evermann (1898). of the Scorpaena except that they lack ally convex; cranial ridges, when pres­ Emmelas Jordan and Evermann (1898), cutaneous filaments, and that their head, ent, low; gill rakers very long and slen­ type species: S. glaucus: No descrip­ less covered with bristles, is scaly. der; scales usually smooth, few if any tion of the subgenus offered; see de­ “There is a large species of them in accessory scales. Parietal ridges ending scription of type species in Jordan and the North Sea, called ‘marulke,’ and in in spines; preocular, supraocular and Evermann (1898). some places ‘carp’ (Sebastes norvegi­ tympanic spines well developed. Peri­ cus, Nob., Perca marina, Pennt., Perca toneum black. Postocular spine pres­ Zalopyr Jordan and Evermann (1898), norvegica, Müll.), Bonnat. Encycl. ent. Second anal spine usually stron­ type species: S. aleutianus: No descrip­ Meth., plate on ichthyology, fig. 210. It ger and longer than third. Symphyseal tion of the subgenus offered; see de­ is red, and is often more than two feet knob strong, projecting forward. Dorsal scription of type species in Jordan and long. It is dried to make food provi­ low. (Peritoneum black, mandibles and Evermann (1898). sions. Its dorsal spines are used as nee­ maxillary scaled).” Sebastiscus Jordan and Starks (1904), dles by Eskimos. Primospina Eigenmann and Beeson type species: Sebastiscus marmoratus: “The Mediterranean has a species (1893), type species: S. mystinus: Taken “This genus is based on species having very similar, but its dorsal rays are less from their key: Parietals meeting above the external appearance of Sebastodes numerous (Sebastes imperialis, Nob., the supra-occipital. Lower jaw much and much resemblance to the subgener­ Scorpaena dactyloptera, Laroche, An­ projecting; head broad, the skull usu­ ic group called Pteropodus, but having nales Mus., 13, plate 22, fig. 9). Its palate ally convex; cranial ridges, when pres­ 12 spines in the dorsal fin and the ver­ is black; it lacks a swim bladder, al­ ent, low; gill rakers very long and slen­ tebrae 10+14=24, agreeing in these re­ though the preceding species has one.”2 der; scales usually smooth, few if any gards with Scorpaena. From Helicole­ Sebastodes Gill (1861), type species: accessory scales. Postocular spine not nus, Sebastiscus differs, solely, perhaps, S. paucispinis: “A very different facies developed. Parietal ridges not ending in the presence of a well-developed air­ from Sebastes, and is readily distin­ in spines. Preocular spines well devel­ bladder. Peritoneum pale.” guished by the longer body, the very oped. Supraocular and tympanic spines Sebastocles Jordan and Hubbs (1925), protuberant lower jaw, which has a sometimes present. Interorbital wide, type species: S. elegans (=S. hubbsi): symphyseal swelling beneath, the very convex. Peritoneum black. Approxi­ “Dorsal spines low, normally fourteen; minute scales, the form and armature of mated edges of sub-opercle, and in­ interorbital deeply concave, size small.” the head, the deep emargination of the teropercle frequently ending in spines. dorsal fin and the emarginated caudal.” Sebastocarus Jordan and Evermann Pteropodus Eigenmann and Beeson (1927), type species: S. serriceps: “This Sebastichthys Gill (1862b), type species: (1893), type species: S. maliger: Taken genus is closely allied to Sebastichthys S. nigrocinctus: “Species referred to the from their key: Parietals separated by from which it differs in the more com­ genus Sebastes, which has eleven to the supra-occipital. Cranium with many pressed body and especially in the strict twelve (XI.+1.–XII.+1.) spines in its first ridges, all ending in spines. Postocular and high ridges on the head, all of them dorsal fin, palatine teeth, and the physi­ spines wanting. Coronal spines none. being free from serrations or accessory ognomy of Sebastes (norvegicus).” Auctospina Eigenmann and Beeson tubercles.” Sebastomus Gill (1864), type species: (1893), type species: S. auriculatus: Taken Sebastopyr Jordan and Evermann S. rosaceus: “Distinguished by the tex­ from their key: Parietals separated by (1927), type species: S. ruberrimus: ture of the bones of the skull, armed or­ the supra-occipital. Cranium with many “This genus is allied to Sebastomus, bital ridges, prefrontals &c.” ridges, all ending in spines. Postocular but of much coarser build, the cranial spines wanting. Coronal spines present. Sebastosomus Gill (1864), type species: spines in the adult being rough with S. melanops: “. . . the genus Sebastich­ Eosebastes Jordan and Evermann blunt spinules.” thys includes at least three genera. The (1896), type species: S. aurora: No de­ Hozukius Matsubara (1934), type spe­ Sebastichthys nigrocinctus is somewhat scription of the subgenus offered; see cies: H. emblemarius: “External: Body related to Scorpaena, and distinguished description of type species in Jordan rather robust and somewhat compressed. by elevated, serrated coronal crests. The and Evermann (1898). Maxillary, and mandible covered with other California species represented by Hispaniscus Jordan and Evermann small ctenoid scales but not on bran­ the Sebastes melanops, seen by me, (1896), type species: S. rubrivinctus: chiostegals. Teeth in villiform bands differ so much that they may be sep­ No description of the subgenus of­ on vomer and palatines. Head much

62(2), 2000 17 spinous; nasal, preocular, supraocular, absent except for the parietal ridges the pointed tips directed backward; pre­ postocular, tympanic, coronal, parietal, which are low but never scaled over; pa­ ocular, postocular, tympanic and pari­ and nuchal spines present; upper and rietals separated; mesethmoid process­ etal spines present; supraocular, coro­ posterior margins of orbit provided with es never directed upward; base of crani­ nal and nuchal spines absent; supra­ many accessory spines; parietal ridge um markedly curved; ventral process of ocular ridges exceedingly high and not conspicuous. Orbital margins of basisphenoid feeble, never attached to stout, forming a narrow flat area be­ preorbital and second suborbital bones the parasphenoid; nasal spines low, but tween them; frontal ridges barely evi­ armed with a sharp spine which is di­ strong; preorbital lobes without spines; dent, can not be seen without elimina­ rected backward; first suborbital mar­ second suborbital bone much wider tion of the skin; parietal ridges broad gined with several small spines; lower than long. Vertebrae usually 30 includ­ and high, but a little lower than the su­ border of preorbital with two very ing the hypural. Peritoneum jet black. praocular ridges; nasal spines stout, di­ strong, sharp bifid spines. Lower three Gill-rakers long and slender. Dorsal rected upward and slightly backward; preopercular spines much larger than mostly XIV, 14; anal usually III, 9 mesethmoid processes strongly com­ upper two; subopercle and interopercle to 10. Lower jaw strongly produced pressed, directed forward and upward; each with a small, sharp spine at their beyond the upper when the mouth is parietal bones separated; base of cra­ approximation. Symphyseal knob very closed; symphyseal knob evident; jaws nium straight; ventral process of ba­ large. Dorsal with 12 spines and about thickly covered by small scales.” sisphenoid entirely meeting the para­ 12 rays; anal with 3 spines and about 6 Mebarus Matsubara (1943), type spe­ sphenoid. Vertebrae 26 including hy­ rays. Auxillary flap absent. cies: S. inermis: “Cranium rather thin; pural. Peritoneum white. Air-bladder “Internal: Skull thick and heavy. interorbital space and occiput plain or entirely free from the layer of connec­ Base of skull (paraspheniod) moder­ very slightly convex, the former about tive tissue lying outside the peritone­ ately curved. Frontal, parietal, pterotic 3.0 times in base of the cranium; pre­ um, the bladder being easily detached and supratemporal much cavernous re­ ocular spines always present; supraoc­ from the coelom; the extrinsic muscle sembling a bee-hive in general appear­ ular and parietal spines small or absent; band of the bladder gives rise poste­ ance. Parietals entirely meeting; upper postocular, tympanic and nuchal spines riorly to a single short ligament; the margin of orbit and frontal ridges some­ entirely absent; cranial ridges absent anterior part of the muscle band not what raised. Mesethmoid processes di­ except for the parietal which is very pierced through by a ligament. Gill­ rected upward and forward. Nodule on low and sometimes scaled over; pa­ rakers very short and blunt. Dorsal the front of prefrontal for the articula­ rietals widely separated; mesethmoid spines 13. Jaws, snout and branchioste­ tion with palatine well developed. Ven­ processes depressed or slightly direct­ gals mostly naked; upper jaw project­ tral process of basisphenoid rudimen­ ed upward; base of cranium strongly ing beyond the lower when mouth is tary, not reaching to paraspheniod; ba­ curved; ventral process of basisphenoid closed; symphyseal knob at the tip of sisphenoid with a small pore between feeble; nasal spines low, but strong. lower jaw absent or barely evident.” it and basis crania. Myodome well de­ Vertebrae usually 26 or 27 including the veloped, with a rather large posterior Neohispaniscus Matsubara (1943), type hypural. Peritoneum black or pale. Gill­ species: S. schlegelii: “Skull thick and opening. Second suborbital bone very rakers long and slender. Dorsal spines broad and long entirely touching to heavy; interorbital space flat or slight­ 13. Lower jaw projecting far beyond ly convex, its width about 2.6 to the upper ridge of preopercle; third the upper, provided with a prominent and fourth suborbital bones present. 3.3 times in base of the cranium; cra­ symphyseal knob; maxillary at least nial spines well developed, directed Gill-rakers long and stout, the longest scaly; preorbital lobes with or without one much longer than the longest gill­ straightly backward; preocular, post­ spines. ocular, tympanic and parietal spines lamella. Vertebrae 9 + 16 + hypural = “The present new subgenus is close­ 26; parapophysis developed from the always present supraocular, corneal and ly related to the subgenus Rosicola, nuchal spines absent; parietal ridges sixth vertebra, its processes widely di­ but differs from it in having weak pari­ verging, directed outward and down­ high and naked; frontal ridges low but etal ridges and flattish interorbital space distinct, the space between them shal­ ward. Air bladder absent. Peritoneum and lacking the postocular and tympan­ jet black.” lowly concave; supraocular edges low ic spines. The subgenus is also separa­ or depressed. as high as or lower than Hatumeus Matsubara (1943), type spe­ ble from Sebastosomus in having usu­ the frontal ridges; mesethmoid process­ cies: S. owstoni: “Cranium thin and ally the strong preocular spine, flattish es slightly elevated upward; base of cra­ papery; interorbital space and occiput interorbital space and parietals widely nium some-what curved; parietals sepa­ flat, the former about 3.3 times in separated.” rated or partly meeting; ventral process base of the cranium; preocular, postoc­ Murasoius Matsubara (1943), type spe­ of the basisphenoid well developed, en­ ular, tympanic and parietal spines pres­ cies: S. pachycephalus: “Cranium very tirely meeting the parasphenoid. Ver­ ent, but small and weak; nuchal spines thick and heavy; interorbital space tebrae 26 including hypural. Peritone­ sometimes present; supraocular and deeply concave, about 31/2 in base of um usually white. Gill-rakers, compar­ coronal spines absent; cranial ridges cranium; cranial spines broad and stout, atively long and slender with pointed

18 Marine Fisheries Review tips. Lower jaw generally projecting directed backward and downward; pre­ Allosebastes Hubbs (1951), type spe­ beyond the upper when the mouth is ocular supraocular, coronal and nuchal cies: S. sinensis: “. . . reduction of the closed, with a rather small symphyseal spines entirely absent; parietal ridges anal soft-rays to 5. . . . smooth, mostly knob. Jaws usually naked. low but broad; supraocular edges de­ cycloid scales; the unswollen lower “The present subgenus is closely re­ pressed, never higher than the frontal pectoral rays; the excessively long lated to Hispaniscus, but differs from it ridges; frontal ridges low, the space be­ anal spines (the second extends well in having a much longer and wider in­ tween them flattish, never deeply con­ beyond the longest soft-ray); the close­ terorbital space, elevated mesethmoid cave; mesethmoid processes directed ly clumped subparallel upper 3 preoper­ processes and shallowly depressed area forward and upward; parietals meeting cular spines (the lower 2 are well sep­ between the frontal ridges.” or narrowly separated; the patch of vo­ arated and divergent); the protuberant merine teeth triangular; base of cranium posterior end of the mandible, almost Takenokius Matsubara (1943), type spe­ straight; ventral process of basisphe­ resembling a flat spine; and, especial­ cies: S. oblongus: “Cranium thick and noid meeting the parasphenoid. Verte­ ly, the very firm well-exposed subor­ heavy; interorbital space and occiput brae 26 including hypural. Peritoneum bital stay (second suborbital), with the flat or slightly convex, the former about pale. Gill-rakers very short, tubercular. bone widened about the unusually large 3.7 times in base of cranium; postocular, Jaws equal in length and entirely scale­ pore, which has a somewhat raised and tympanic and parietal spines present, less; symphyseal knob inconspicuous.” roughened rim.”

62(2), 2000 19 Appendix II Summary of Nomenclature of Sebastes Listed Alphabetically by Subgenus

Eigenmann and Beeson, Original species1 Authors Date Present species2 Common name 1894

(Fitch) 1964 phillipsi chameleon rockfish Lea and Fitch 1979 melanosema semaphore rockfish (Westrheim and Tsuyuki) 1967 reedi yellowmouth rockfish Barsukov 1988 cheni (Taranetz and Moiseev) 1933 polyspinis northern rockfish Kim and Lee 1994 koreanus hwanghee-bolnak Eitner et al. 1999 moseri whitespeckled rockfish alutum (Gilbert) 1890 alutus Pacific ocean perch Acutomentum brevispine (Bean) 1884 brevispinis silvergray rockfish Pteropodus clavilatum Starks 1911 (?) eigenmanni Cramer 1896 (?) entomelas (Jordan and Gilbert) 1880 entomelas widow rockfish Primospina (Cramer) 1895 hopkinsi squarespot rockfish (Eigenmann and Beeson) 1893 macdonaldi Mexican rockfish Acutomentum ovale (Ayres) 1863 ovalis speckled rockfish Acutomentum rufum (Eigenmann and Eigenmann) 1890 rufus bank rockfish Sebastomus (Jordan and Starks) 1904 flammeus sankou-menuke (Jordan and Starks) 1904 iracundus oo-saga (Jordan and Snyder) 1900 scythropus ukeguchi-mebaru (Wakiya) 1917 baramenuke bara-menuke paucispinosus Matsubara 1943 (alutus)

Chen 1975 varispinis Chen 1975 peduncularis (Beebe and Tee-Van) 1938 cortezi (Gilbert) 1890 sinensis Pteropodus Quast 1971 variegatus harlequin rockfish Lea and Fitch 1972 rufinanus dwarf-red rockfish (Gilbert) 1890 diploproa splitnose rockfish Sebastichthys (Starks) 1911 emphaeus Puget Sound rockfish prorigerum (Jordan and Gilbert) 1880 proriger redstripe rockfish Pteropodus (Gilbert) 1890 saxicola stripetail rockfish Pteropodus (Gilbert) 1897 semicinctus halfbanded rockfish (Gilbert) 1915 wilsoni pygmy rockfish (Gilbert) 1890 zacentrus sharpchin rockfish Pteropodus dallii (Eigenmann and Beeson) 1894 dalli calico rockfish Pteropodus auriculata Girard 1854 auriculatus brown rockfish Auctospina

Hilgendorf 1880 glaucus kuro-menuke

(Gilbert) 1890 aurora aurora rockfish Auctospina (Jordan) 1897 crameri darkblotched rockfish deani Starks 1911 (?) introniger (Gilbert) 1890 (melanostomus) Acutomentum (Eigenmann and Eigenmann) 1890 melanostomus blackgill rockfish Acutomentum rupestris (Gilbert) 1890 (melanostomus) Sebastomus

(Jordan and Thompson) 1914 owstoni hatsume

Ayres 1859 elongatus greenstriped rockfish (Eigenmann and Eigenmann) 1889 levis cowcod Sebastomus (Jordan and Gilbert) 1880 rubivinctus flag rockfish Sebastichthys atrorubens (Gilbert) 1898 (atrovirens) (Jordan and Gilbert) 1880 atrovirens kelp rockfish Pteropodus (Jordan and Hubbs) 1925 thompsoni usu-mebaru (Matsubara) 1934 wakiyai gaya-modoki Cuvier and Valenciennes 1829 inermis mebaru Gunther 1878 joyneri togotto-mebaru taczanowskii (Steindachner) 1880 taczanowski ezo-mebaru Barsukov 1972 minor aka-gaya guentheri (Jordan and Starks) 1904 (inermis) paradoxus Matsubara 1943 (wakiyai) tokionis (Jordan and Starks) 1904 (inermis)

Matsubara 1943 *nudus Temminck and Schlegel 1843 pachycephalus mura-soi (Schmidt) 1931 *nigricans Matsubara 1943 *chalcogrammus

Hilgendorf 1880 schlegeli kuro-soi Steindachner and Doderlein 1884 vulpes kitsune-mebaru

20 Marine Fisheries Review Major generic or subgeneric assignments Other subgeneric assignments Jordan and Evermann, Jordan, Evermann, Current 1898 and Clark, 1930 Matsubara, 1943 Reference Subgenus subgenus

Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown Unknown

Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Chen, 1986 Acutomentum Acutomentum Acutomentum Acutomentum Chen, 1986 Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Chen, 1986 Acutomentum Acutomentum Acutomentum Acutomentum Chen, 1986 Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum Acutomentum

Chen, 1986 Allosebastes Allosebastes Chen, 1986 Allosebastes Allosebastes Chen, 1986 Allosebastes Allosebastes Hispaniscus Hispaniscus Hubbs, 1951; Chen, 1986 Allosebastes Allosebastes Chen, 1986 Allosebastes Allosebastes Allosebastes Eosebastes Eosebastes Chen, 1986 Allosebastes Allosebastes Eosebastes Chen, 1986 Allosebastes Allosebastes Acutomentum Acutomentum Chen, 1986 Allosebastes Allosebastes Eosebastes Eosebastes Chen, 1986 Allosebastes Allosebastes Eosebastes Eosebastes Chen, 1986 Allosebastes Allosebastes Acutomentum Chen, 1986 Allosebastes Allosebastes Hispaniscus Hispaniscus Chen, 1986 Allosebastes Allosebastes

Auctospina Auctospina Auctospina Auctospina Auctospina

Emmelas Emmelas Emmelas Emmelas

Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Eosebastes Sebastomus Sebastomus Eosebastes

Hatumeus Hatumeus

Hispaniscus Hispaniscus Hispaniscus Hispaniscus Hispaniscus Hispaniscus Hispaniscus Hispaniscus Hispaniscus Zalopyr Zalopyr Mebarus Zalopyr Zalopyr Chen, 1985; 1986 Mebarus Mebarus Mebarus Chen, 1985; 1986 Mebarus Mebarus Mebarus Chen, 1985 not Mebarus Mebarus Mebarus Chen, 1985; 1986 Mebarus Mebarus Mebarus Chen, 1985; 1986 Mebarus Mebarus Mebarus Chen, 1985 not Mebarus Mebarus Mebarus Mebarus Mebarus Mebarus Mebarus Mebarus Mebarus

Murasoius Murasoius Murasoius Murasoius Murasoius Murasoius Murasoius Murasoius

Neohispaniscus Neohispaniscus Neohispaniscus Chen, 1986 Takenokius Neohispaniscus

continued

62(2), 2000 21 Appendix II (continued)

Eigenmann and Beeson, Original species1 Authors Date Present species2 Common name 1894 ijimae (Jordan and Metz) 1913 (vulpes) zonatus Chen and Barsukov 1976 (vulpes)

(Jordan and Gilbert) 1880 carnatus gopher rockfish Pteropodus Richardson 1844 caurinus copper rockfish Pteropodus (Jordan and Gilbert) 1881 chrysomelas black-and-yellow rockfish Pteropodus gilberti Cramer 1896 (dalli) (Jordan and Gilbert) 1880 maliger quillback rockfish Pteropodus Ayres 1854 nebulosus China rockfish Pteropodus (Jordan and Gilbert) 1880 rastrelliger grass rockfish Pteropodus vexillaris (Jordan and Gilbert) 1880 (caurinus) Pteropodus Hilgendorf 1880 nivosus goma-soi Hilgendorf 1880 trivittatus shima-zoi (Matsubara) 1937 hubbsi yoroi-mebaru (Matsubara) 1934 longispinis kourai-yoroi mebaru alexandri (Evermann and Goldsborough) 1907 (auriculatus) (Thompson) 1915 babcocki redbanded rockfish (Jordan and Gilbert) 1880 miniatus vermilion rockfish Sebastomus (Gill) 1864 pinniger canary rockfish Sebastomus

(Travin) 1951 mentella deepwater redfish Storer 1854 fasciatus Acadian redfish norvegica (Ascanius) 1772 norvegicus golden redfish Kroyer 1845 viviparus Norway haddock

Ayres 1859 nigrocinctus tiger rockfish Sebastichthys albo-fasciatus • (Lacepede) 1802 albofasciatus ayame-kasago (Cuvier and Valenciennes) 1829 marmoratus kasago tertius (Barsukov and Chen) 1978 (marmoratus)

(Jordan and Gilbert) 1880 serriceps treefish Sebastichthys

(Eigenmann and Eigenmann) 1890 goodei chilipepper Sebastodes (Gilbert) 1896 jordani shortbelly rockfish Ayres 1854 paucispinis bocaccio Sebastodes (Jordan and Starks) 1904 itinus yanagi-mebaru Hilgendorf 1880 steindachneri yanagi-no-mai

Chen 1975 spinorbis Chen 1971 exsul Chen 1971 simulator pinkrose rockfish Chen 1971 rosenblatti greenblotched rockfish Chen 1971 ensifer swordspine rockfish Chen 1971 notius Ayres 1859 helvomaculatus rosethorn rockfish Sebastomus Chen 1971 lentiginosus freckled rockfish ayresii (Gilbert and Cramer) 1897 (rosaceus) (Jordan and Gilbert) 1880 chlorostictus greenspotted rockfish Sebastomus (Jordan and Gilbert) 1880 constellatus starry rockfish Sebastomus (Eigenmann and Eigenmann) 1890 eos pink rockfish Sebastomus gillii (Eigenmann) 1891 gilli bronzespotted rockfish Sebastomus rhodochloris (Jordan and Gilbert) 1880 (helvomaculatus) Sebastomus Girard 1854 rosaceus rosy rockfish Sebastomus (Jordan and Gilbert) 1882 umbrosus honeycomb rockfish Sebastomus Valenciennes 1833 oculatus (Gmelin) 1789 capensis chamaco (Evermann and Radcliffe) 1917 (capensis)

(Tilesius) 1813 ciliatus dusky rockfish Sebastosomus (Ayres) 1862 flavidus yellowtail rockfish Sebastosomus Girard 1856 melanops black rockfish Sebastosomus (Jordan and Gilbert) 1881 mystinus blue rockfish Primospina (Eigenmann and Eigenmann) 1890 serranoides olive rockfish Sebastosomus

(Jordan and Gilbert) 1883 ruberrimus yelloweye rockfish

Gunther 1877 oblongus takenoko-mebaru

Barsukov 1970 borealis shortraker rockfish (Jordan and Evermann) 1898 aleutianus rougheye rockfish kawaradai (Matsubara) 1934 (aleutianus) melanostictus (Matsubara) 1934 (aleutianus) swifti (Evermann and Goldsborough) 1906 (aleutianus) Hilgendorf 1880 matsubarae akou-dai

1 Listed when different from present species name. 2 ( )=synonomyzed with; *=form of pachycephalus.

22 Marine Fisheries Review Major generic or subgeneric assignments Other subgeneric assignments Jordan and Evermann, Jordan, Evermann, Current 1898 and Clark, 1930 Matsubara, 1943 Reference Subgenus subgenus

Neohispaniscus Chen, 1986 Takenokius Neohispaniscus Chen, 1986 Takenokius Neohispaniscus

Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Pteropodus Chen, 1986 Takenokius Pteropodus Pteropodus Chen, 1986 Takenokius Pteropodus Pteropodus Jordan and Hubbs, 1925 Sebastocles Pteropodus Pteropodus Pteropodus

Rosicola Rosicola Rosicola Rosicola Rosicola Rosicola Rosicola Rosicola Rosicola Rosicola

Sebastes Sebastes Sebastes Sebastes Sebastes Sebastes

Sebastichthys Sebastichthys Sebastichthys

Matsubara, 1943 Sebastiscus Sebastiscus Matsubara, 1943 Sebastiscus Sebastiscus Chen, 1986 Sebastiscus Sebastiscus

Sebastichthys Sebastocarus Sebastocarus

Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes Sebastodes

Chen, 1975 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus Chen, 1971 Sebastomus Sebastomus

Primospina Sebastosomus Sebastosomus Sebastosomus Sebastosomus Sebastosomus Sebastosomus Sebastosomus Sebastosomus Primospina Sebastosomus Sebastosomus Sebastosomus Sebastosomus Sebastosomus

Sebastomus Sebastopyr Sebastropyr

Takenokius Chen, 1986 Takenokius Takenokius

Zalopyr Zalopyr Zalopyr Zalopyr Zalopyr Zalopyr Zalopyr Zalopyr Eosebastes Zalopyr Zalopyr Zalopyr

62(2), 2000 23