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An historical review of Sebastes Taxonomy and Systematics.

Discovery: The Late 1700's

The systematics of the speciose genus Sebastes (1), rockfishes, particularly in the North Pacific, have challenged ichthyologists and others even casually interested in these fishes for well over a century. Fernholm and Wheeler (1983) detailed problems associated with the early descriptions of Sebastes. The first scholarly reference to a rockfish was by Linnaeus (1761) who included Norway and Italy as the range of a fish he described in 1758 and named Perca marina. In his former publication (Linnaeus, 1758), he also described Perca scriba, a Mediterranean serranid. Reiterating his description of Perca marina, Linnaeus (1761) cited the common Norwegian name (uer or rodfisk) for the fish from Norway, making it clear that he was including a rockfish, as well as a fish from the Mediterranean region in his description.

Cuvier (1829) was the first to use the generic name Sebastes, and in his second, and much more thorough description of the genus (Cuvier and Valenciennes, 1829) he included descriptions of species from the North Atlantic (Sebastes norvegicus), the Mediterranean Sea (S. imperialis = Helicolenus dactylopterus), and the Southern Hemisphere (S. capensis). (2) Cuvier and Valenciennes (1829) recognized the confusion between the northern fish and Perca marina, which he said was "hardly believable"; nevertheless, the name Sebastes marinus (Linnaeus) was used until recently for this fish from the North Atlantic. However, Fernholm and Wheeler (1983) found that the specimen Linnaeus used as the basis of his description was in fact the Mediterranean serranid Serranus scriba, so Sebastes (Perca) marina became a synonym of Serranus scriba, and the rockfish was left without a type species for reference. This made way for recognizing Ascanius (1772) as the author of Perca norvegica (i.e. Sebastes norvegicus), the most common rockfish of the northeastern Atlantic.

Proliferation of Subgenera: 1861-1898

By 1845, similar fish had been described as species of Sebastes from both the eastern and western North Pacific. In 1854, Ayres discovered Sebastes paucispinis from off California (Ayres, 1854a). Since Sebastes paucispinis is very different from Sebastes norvegicus, but clearly related to it, its discovery prompted Gill (1861) to erect another genus for it: Sebastodes. This began a trend by Gill and others to establish generic or subgeneric groupings as more and more similar species were described on both sides of the North Pacific during the 1860's-1880's (Fig. 1, 2). By 1898, Jordan and Evermann grouped the 55 northeast Pacific species that they recognized into 13 subgenera (Jordan and Evermann, 1898). They also mentioned 13 species from Japan and 3 species from the Southern Hemisphere. Generally the characters of a proposed subgenus were only those of the type species for the subgenus. In most cases the definitions of the subgenera were inferred rather than explicitly stated (Appendix I).

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Confusion: 1863-1900

There was considerable disagreement among ichthyologists of the late 1800's concerning these subgenera of northeast Pacific rockfishes and the importance of the characters used to distinguish them. Reading the papers of Ayres (1854a, b, 1855, 1859, 1863a, b, c, d), Gill (1861, 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 (Sebastodes 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 species 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 included with the North Atlantic species in Sebastes. In Gill's 1864 paper "Critical remarks on the genera Sebastes and Sebastodes of Ayres" he stated: "The value of the characters used to distinguish the genera Sebastes, Sebastichthys and Sebastodes are now indeed so generally conceded by scientific men, that it is unnecessary to further argue in their favor. I shall only remark that the combinations and distinction of forms by Dr. Ayres are alike unnatural and violate all natural affinities ..." In discussing Ayres' ideas on affinities of flat-fishes, Gill (1864) stated: "Dr. Ayres ... ideas of affinity are extremely crude and unreliable, nothing can be learned from them." However, in discussing this paper Gunther (1865) commented: "Dr. Gill would advance ichthyology by giving us serviceable descriptions, instead of limiting himself to synoptical tables with minute sub-generic subdivisions. As regards his frequent critical remarks on synonyms, it would be very useful if he would state whether he arrived at his conclusions from an examination of typical specimens, but it is not even evident whether he has known the species from autopsy." Ayres (Fig. 3) was not the only naturalist from San Francisco to receive severe criticism from Gill (Fig. 4): "Of course, Gill had an agenda for this criticism for he objected to the intrusion of these Californian upstarts, amateurs in his opinion, in what he had carved out as his personal fiefdom, the fishes of the North Pacific" (Leviton and Aldrich, 1997, footnote 16.3, p. 196).

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In 1880, Jordan and Gilbert (1880) described 7 new species from California, and after trying to place them in the four genera then recognized by Gill, they left paucispinis in Sebastodes and stated that: "the groups Sebastosomus and Sebastomus cannot be maintained as genera distinct from Sebastichthys, and that, in order to recognize them as subgenera even, a different distribution of species must be adopted."

Eigenmann and Beeson (1893, 1894) tried to bring some order to the argument by publishing a key and diagram of relationships for the eight subgenera they recognized (with mystinus as the sole member of Primospina representing the base from which all other species radiate) (Fig. 5). However, a footnote by the editor (R. Edward Earll) to the title of the 1894 paper which appeared in the Proceedings of the U.S. National Museum stated: "The classification adopted by the authors of this paper is based on their own peculiar interpretation of the importance of certain structural characters. The arrangement and nomenclature proposed here will not be, at present at least, followed by the National Museum." Further, Cramer (1895), in his detailed study of the cranial osteology of these fishes, took strong exception with the importance of the first dichotomy (union or nonunion of the parietals) in the key of Eigenmann and Beeson (1893). Since Cramer's work was in essence a rebuttal of the work of Eigenmann and Beeson, he included their entire article as an appendix to his paper. Cramer (1895) did not explicitly recognize any subgenera for the 32 species of Sebastes he examined, but his key to species based on cranial characters exactly groups the species into the subgenera eventually used by Jordan and Evermann (1898) in their classic "The Fishes of North and Middle America." In the introduction to their species accounts of Sebastes, Jordan and Evermann (1898) included from Cramer (1895) extensive quotations from his text, and his entire key to species, which they amended slightly by adding a few species Cramer did not consider, and from Eigenmann and Beeson (1893) their key to subgenera. Jordan and Evermann (1898) commented on Eigenmann and Beeson's (1894) work saying: "Messrs. Eigenmann & Beeson have attempted to subdivide this genus into several subgenera on the basis of cranial characters. Mr. 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 contact (not union) of the parietals, seems to us of very slight value, even at a specific distinction."

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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 Evermann (1927), and one by Hubbs (1951). Through the first half of the 20th century, authors variously lumped, split, or ignored subgenera completely. Although in their "Checklist of Fishes of North and Middle America," Jordan et al. (1930) placed 66 species of rockfishes they recognized in 16 genera (that is, they elevated the subgeneric names to generic status), most American researchers used only two genera: Sebastes for Atlantic species and Sebastodes for Pacific species, disregarding the subgenera.

Matsubara (1943) (Fig. 6) presented a very detailed analysis of the Scorpaenidae of the Northwest Pacific. He subdivided the family into 14 subfamilies, and in the Sebastinae he included four genera, one of which was Sebastes. He grouped the 30 species of Sebastes he considered into 10 subgenera, 5 of which were newly erected. The five previously established subgenera he used had representative species in the Northeast Pacific also. Matsubara (1943) provided a key to the subgenera he used, and descriptions based on a number of osteological, meristic, and morphometric characters. He also concluded that there was no validity in grouping the Atlantic species of Sebastes separately from the Pacific species (then considered in Sebastodes), and since Sebastes is the older name it takes precedence.

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The early-described Pacific species have fewer dorsal spines and vertebrae than the Atlantic species, characters thought important enough to justify placing the species from the two oceans in separate genera. However, species were later found in the northern Pacific Ocean and Bering Sea that have vertebral and dorsal spine counts approaching those of the Atlantic species, calling into question the rationale for maintaining the generic distinction. This change in thinking is reflected in successive editions of the American Fisheries Society's "List of Common and Scientific Names of Fishes from the United States and Canada." The first two editions (AFS, 1948; Bailey et al., 1960) recommended Sebastes for Atlantic species and Sebastodes for Pacific species; however, later editions (Bailey et al., 1970; Robbins et al., 1980; 1991) recommended using Sebastes for rockfishes from both the Atlantic and Pacific waters of North America.

Limited Morphological Studies: the 1970's

Chen (1971) reviewed Sebastomus, the most distinctive subgenus of Sebastes. Occurring in the northeast Pacific and in the Southern Hemisphere, Sebastomus contains 16 species, which are all characterized by having strong head spines, the second anal spine longer than the third, and a series of six large white blotches (one at the tip of the opercle and five between the dorsal fin and lateral line [not all are present in all species]). Chen (1971) gave detailed accounts of the species and constructed a diagram of relationships among the 13 then-known species based on a subjective analysis of about 15 characters. He considered phenetic similarity among the species in constructing his diagram. The characters he used were color patterns, body configuration (morphometrics and head spines), and meristics, He did not specify the out-group for his analysis, or what he considered to be the primitive states of the characters he used. Without this information and with the informal discussion of relationships he offered, it is difficult to evaluate his conclusions. Thus, even within this relatively well-known and cohesive species group, considerable additional work is needed.

Hallacher (1974) characterized the structure and points of attachment of swimbladder muscles for 82 species of Sebastes, and found two major types and seven patterns within these types. In general, he found little correspondence between species sharing the same muscle patterns and their placement in the proposed subgenera (including Sebastomus). For example, two similar appearing members of Sebastodes (S. brevispinis and S. paucispinis) had major differences in structure. However, all but one species of Pteropodus shared the same swimbladder muscle pattern.

During this period several new species were described or reviewed (e.g. Barsukov (1970), Chen (1975), Tsuyuki and Westrheim (1970)), and some studies dealt with the component species of some subgenera (e.g. the Sebastes vulpes "complex": Chen and Barsukov (1976), Sebastiscus: Barsukov and Chen (1978), and Mebarus: Chen (1985)).

Systematic morphological analysis of the entire genus during this period was limited to Barsukov (1981), who published a complex diagram of inter-specific relationships of the Sebastinae based on a dubious speciation hypothesis (Kendall, 1991). Barsukov (1981) proposed that there were seven cycles in the evolutionary history of the Sebastinae. "In each one no more than three descendant species originated sympatrically from one ancestral species. Such a triad is characteristic of speciation with incomplete (mainly biotopic) isolation. A species, once established at intermediate depths, emerges into the extreme position of another dimension, i.e., it develops ecological and morphological features unique for the trio, and usually associated with adaptations to life in open water" (Barsukov, 1981:1). Based on this premise he reviewed the sequence of geological events that led to the divergence found in some lineages of these fishes, and related the morphology of living forms to their history, ecology, and zoogeography. For example, in the first cycle he considered the subfamily Sebastinae to be composed of three genera: the shallow water Helicolenus with 13 species, the intermediate depth Sebastes with 102 species, and the deep-water Hozukius with 2 species. In the second cycle he felt that the genus Sebastes split into three subgenera: the shallow water Sebastiscus with 3 species, the intermediate depth Sebastodes with 88 species, and the deep-water Sebastes with 14 species. More divergence occurred in the intermediate-depth branch than in the other two during each cycle, and some branching was not successful (incipient species became extinct). Using these questionable evolutionary principles Barsukov (1981) diagramed the relationships of all known species of the Sebastinae.

Genetics Enters the Picture: the Late 1900's

Starting in the 1960's, newly developed genetic techniques were applied to systematic studies of rockfishes. Barrett et al. (1966) examined electrophoretic patterns of hemoglobin in 27 species of Sebastes from off California and found that each species, including some morphologically similar species pairs (e.g. Sebastes caurinus-S, vexillaris, S. eos-S, chlorostictus), had a distinctive pattern. They also found shared patterns among species in some subgenera (e.g. Pteropodus, Sebastomus), but distinct differences among some members of other subgenera (e.g. Sebastosomus). This work was closely followed by a study (Tsuyuki et al., 1968) examining hemoglobin, eye lens, and muscle protein electrophoretic patterns in numerous species of Sebastes from both the Atlantic and Pacific, as well as representatives of other scorpaeniform genera. As in Barrett et al. (1966), species-specific patterns were found in hemoglobins, and they interpreted within-species variation as evidence of additional undescribed species. Muscle proteins were not all species-specific, but they separated the 27 species of Pacific Sebastes they examined into four subgroups. Comparing these subgroups with the subgenera, little correspondence is seen. Among the 27 species, 11 subgenera are represented. Sebastes aurora, the sole member of Eosebastes, had a distinct pattern. As in Barrett et al. (1966), there was some coherence among the species of Pteropodus; two species (S. caurinus and S. nebulosus) grouped together as distinct from the other species. However, another species of Pteropodus (S. maliger) was in a group of 12 species, representing 8 subgenera in which there were no relationships among the species. The muscle and eye lens proteins were distinct at the generic level.

In the early 1970's, Johnson et al. (1970a, b, 1971, 1972) and Johnson (1972) conducted a series of investigations on intraspecific variation in Sebastes alutus and other species of Sebastes, interspecific variation among Sebastes, and intergeneric variation within the Scorpaenidae, using electrophoresis of muscle proteins and several enzymatic systems. Polymorphisms were found in 11 of the 31 species studied. Three patterns were seen in S. alutus using two enzyme systems, however these patterns were not related to the geographic distribution of the samples. Among 27 species of Sebastes from the Pacific, 10 had unique patterns that allowed them to be identified based on the muscle protein and enzyme systems used in these studies. Several species demonstrated differences in only one enzyme system and thus appeared to be closely related to each other (e.g. S. reedi and S. crameri and S. caurinus, S. auriculatus and S. maliger). There was little variation between Atlantic and Pacific Sebastes, which were clearly distinct from Sebastolobus and Helicolenus.

Wishard et al. (1980) used protein electrophoresis to examine within-species relationships among five species of Sebastes from the northeast Pacific. Based on allele frequencies at 21 loci, they found evidence of three populations of S. alutus, two of S. pinniger, and one each of S. flavidus, S. goodei, and S. paucispinis.

At about the same time, genetic methods were being used in studies on the morphologically similar North Atlantic species of Sebastes (Nefyodov, 1971; Naevdal, 1978; Payne and Ni, 1982). A major focus of these studies using electrophoresis of hemoglobin and various enzyme systems was to confirm the presence of more than one sympatric species (S. mentella, S. fasciatus, and S. norvegicus). More recent work on Atlantic rockfish has employed genetic techniques to identify specimens, particularly juveniles, where more than one species occurs (Nedreaas and Naevdal, 1991; Rubec et al., 1991), and to examine population structure within S. norvegicus (Nedreaas et al., 1994).

Electrophoretic studies on Pacific Sebastes reached a plateau with the work of Seeb (1986). She analyzed electrophoretic variations of 28 enzymes in 48 species of Sebastes and was able to identify all but a few closely related species pairs on the basis of this analysis. She produced phenograms based on genetic distances among the species. As in previous studies (see above), species in some subgenera grouped closely together in these diagrams, while some of the branches contained members of several subgenera. For example, all the members of Zalopyr, Sebastes, and Sebastomus that she considered grouped together, as did most species of Pteropodus, and Sebastodes. However, species in Acutomentum, Allosebastes, and Sebastosomus were scattered among several lineages within her diagrams. Further work with allozymes of Pacific rockfish investigated population structure within S. alutus (Seeb and Gunderson, 1988: little structure was found), and the applicability of allozymes in identifying larval and juvenile specimens (Seeb and Kendall, 1991: allozyme activity and resolution decreased with decreasing size of the fish, but was adequate to aid in identification of juveniles and larvae).

In the 1990's, newer, more powerful genetic techniques were applied to rockfish taxonomic studies in both the Atlantic and northeast Pacific. McGauley and Mulligan (1995) amplified the mitochondrial rRNA (mtDNA) genes of Sebastes flavidus using polymerase chain reaction (PCR) and then used restriction fragment length polymorphism (RFLP) analysis to examine population structure. They found essentially no variation in haplotype frequencies among fish collected from Vancouver Island (British Columbia), Westport (Washington), and Cordell Bank (California), indicating that gene flow may be unrestricted within this species throughout its range. They attributed this to long-range dispersal of larvae. However, analysis of a more variable region of the mtDNA may have shown differences that their analysis failed to find. Seeb (1998) investigated gene flow among S. auriculatus (3), S. caurinus, and S. maliger using both allozymes and restriction analysis of mtDNA and found evidence of introgression between all three species in fish from Puget Sound, indicating some level of hybridization.

A major step forward in genetic studies occurred as Rocha-Olivares with others (Rocha-Olivares, 1998a; Rocha-Olivares et al., 1999a, b) began determining the sequence of nucleotides in the cytochrome b gene and part of the control region of the mtDNA molecule (a total of 1,633 bases) in Pacific Sebastes, particularly with members of the subgenus Sebastomus. These data were then used to address a number of taxonomic and systematic questions. An examination of 14 species of Sebastomus, and 40 other species of Sebastes, showed that the subgenus was monophyletic. Low levels of divergence in the genetic data indicated recent rapid radiation of the subgenus within the last million years (Fig. 7). Among Sebastomus, S. rosaceus appeared to represent the oldest lineage, and the rest of the species belonged to one of two clades: a northern clade and a southern clade. Generally, the species pairs produced by the molecular data were the same ones that Chen (1971) found based on morphological data.

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Rocha-Olivares (1998b) used multiplex haplotype-specific PCR primers to create subgenus and species-specific assays for portions of the cytochrome-b gene in four species in the subgenus Sebastomus. He then screened 250 adults representing 31 species of Sebastes that had been identified morphologically, and found that with few exceptions, specimens were correctly assigned to species using the primers. Larvae were correctly assigned to the subgenus Sebastomus, although none was identified as a species from which the primers were developed. Rocha-Olivares and Vetter (1999) also used sequence data to examine intraspecific variation within Sebastes helvomaculatus collected at five sites from Fairweather Bank in the Gulf of Alaska to Cordell Bank off California. They found a high degree of population genetic diversity which could be related to their life history and zoogeography.

The application of genetic techniques is well demonstrated by their resolution of taxonomic problems associated with Sebastes from the Southern Hemisphere. Sebastes of the Southern Hemisphere has proven to be an exceptional challenge ever since S. capensis was placed in the genus by Cuvier and Valenciennes (1829). By the time of Chen's review of Sebastomus (Chen, 1971), 11 nominal species of Sebastes had been described from the Southern Hemisphere, all apparently in Sebastomus. After reviewing the literature (e.g. De Buen, 1960), and specimens from the west coast of South America and from South Africa, Chen (1971) concluded that more work was needed, but that he would consider all nominal species to be forms of S. capensis, as have others (Norman, 1937; Kong Urbina, 1985).

However, Eschmeyer and Hureau (1971) considered the Sebastes off the west coast of South America (S. oculatus) different from those around south central Atlantic islands and off South Africa (S. capensis), and suggested that more than one species occurs off the Pacific coast of South America. Chen (1971) and Eschmeyer and Hureau (1971) both pointed to the limited differentiation of Sebastes below the equator as evidence of the difficulty for this Northern Hemisphere, temperate and subarctic genus to cross the tropics.

Based on genetic sequence data, Rocha-Olivares et al. (1999b) found that within the southern clade of Sebastomus, S. constellatus represented the stock that gave rise to the Southern Hemisphere species: S. oculatus off South America and S. capensis off Africa. Further genetic studies (Rocha-Olivares et al., 1999c) on Southern Hemisphere Sebastes demonstrated that the tropical barrier was crossed successfully only once, and that three lineages (cryptic species?) now exist. The ancestral species is represented by S. capensis which occurs off Tristan da Cuhna and South Africa; this lineage is also present in the southwest Atlantic. There are two lineages of S. oculatus off the coasts of South America: one off the Pacific coast and the other off the Atlantic coast.

Recent work on the Atlantic species of Sebastes has used advanced genetic techniques to investigate relationships among the three morphologically similar sympatric species (e.g. Bentzen et al., 1998; Roques et al., 1999). Bentzen et al. (1998) found an unusual number of tandemly repeated copies of a section of the mtDNA molecule in all three species. The number of tandem repeats (9-17, mean 12.43) was similar in all three species, suggesting that the duplication might have preceded the divergence of the species. They speculated that the Pacific sister species of the Atlantic species might have homologous repeats. They investigated this hypothesis with S. aleutianus, which is another member of the subgenus Sebastes according to Barsukov (1981), but did not find the tandem repeats. They suggested that other Pacific species could be examined to determine if they had the tandem repeats and thus might represent the lineage from which the Atlantic species arose. Roques et al. (1999) found that microsatellite data from eight loci could be used to assign individual fish to the correct species more precisely than with other genetic techniques. It was less likely to assign individuals to the correct population within species. They also concluded, on the basis of shared alleles, that S. norvegicus represents the lineage from which the other Atlantic species arose, and that S. fasciatus and S. viviparus are more closely related to each other than to the other Atlantic species.

As new genetic techniques are developed, they continue to be applied to systematic and taxonomic studies of rockfishes at a variety of levels. For example, Johns and Avise (1998) sequenced the mitochondrial cytochrome b gene from 28 species of rockfish and used these data, as well as the allozyme data of Seeb (1986), to construct molecular phylogenies. They then compared these phylogenies with those of other groups of fishes (Lake Victoria cichlids and icefishes) in which "species flocks" occur. They found that many of the speciation events in Sebastes occurred in clusters in time, and that these events were far more ancient than similar events in African cichlids. They concluded that rockfishes are an ancient species flock, with significant radiation occurring about 5 million years ago (3.6-18 million years ago depending on assumptions concerning rates of evolution).

With the rapid advances in genetic techniques that are now occurring, it is difficult to predict the limits of what will be known of rockfish taxonomy and systematics just a few years from now. However, as more genomic data are accumulated from these phylogenetically interesting fishes, further advances in our understanding of relationships among them and their evolution are assured.

Present Views on the Subgenera

Although the subgeneric names are mentioned occasionally in recent literature, all but Sebastomus still lack rigorous definitions. There is also a lack of agreement concerning the appropriate subgenus for some species (Appendix II). Some species have been assigned to several different subgenera by various authors through the years. For example, Allosebastes was erected for Sebastes sinensis alone by Hubbs (1951); however, Chen (1986) placed 11 other species in the subgenus, without giving a reason for his action.

Based on the literature, subgeneric names have been assigned to 96 of the currently recognized species of Sebastes (Table 1). There are 23 subgenera containing from 1 to 16 species. Nine subgenera contain only one species. Five subgenera have species in both the northwest and northeast Pacific Ocean, six have species only in the northwest Pacific, and ten are exclusively in the northeast Pacific. The North Atlantic contains a subgenus, Sebastes, that occurs only there, while the species in the Southern Hemisphere are in Sebastomus, which also occurs in the northeast Pacific. Some of the subgenera have been proposed as genera, and some groups of subgenera have been considered as genera. No rigorous analysis of the validity of the subgenera has yet been attempted, except for Sebastomus (Rocha-Olivares, 1999a).

Several field guides and keys for northeast Pacific Sebastes have been published (Phillips, 1957; Hitz, 1965; Miller and Lea, 1972; Kramer and O'Connell, 1988; Orr et al., 2000) which assist greatly in identifying the sometimes bewildering array of rockfishes caught even at a single location (Table 2). These guides generally rely on head spine, meristic, morphometric, and color characters that are probably not indicative of systematic relationships among species. The subgenera are usually not mentioned in field guides, although Sebastomus is occasionally noted since the species are quite distinctive externally.

Ishida (1984) reviewed what he considered to be the Japanese Sebastinae (Sebastes, 26 species; Helicolenus, 3 species; Hozukius, 2 species; and Sebastiscus, 2 species). He included a careful evaluation of all species in these genera that occur around Japan. He summarized his views on the relationships within Sebastes, saying "... Sebastes and Sebastiscus of Matsubara (1943) are adopted provisionally and subgenera are not used until world-wide review of these fishes is carded out."

Challenges

Where Does Sebastes Belong?

Sebastes is very provisionally placed in the order Scorpaeniformes, suborder Scorpaenoidei, family Sebastidae, subfamily Sebastinae (Nelson, 1994). Much work is yet to be done on the systematics of Sebastes, beginning at the ordinal level (is the Scorpaeniformes monophyletic? How does it relate to the Perciformes?). The placement of Sebastes within the Scorpaeniformes is presently uncertain. For example, if the Perciformes and Scorpaeniformes had a common ancestor, could it have been similar to Sebastes? Sebastes and basal perciforms share many characteristics. The hallmark of the scorpaeniforms, the suborbital stay is much reduced in Sebastes, and, except for the suborbital stay, Sebastes is more similar to basal perciforms than are other scorpaeniforms. In other ways, Sebastes is the least specialized of the scorpaeniforms: e.g. head spination is minimal, squamation is normal. Is this due to convergence or a common ancestor? Basic unresolved questions regarding the origin and relationships of Sebastes and its close relatives include: was their ancestor tropical or boreal, and was it demersal or pelagic? Does Sebastes represent the point of divergence that led the demersal-boreal scorpaeniforms away from the pelagic-tropical perciforms, or does the genus represent an end point in scorpaeniform evolution (Washington et al., 1984a)? Although these questions may eventually be resolved using cladistic methodology, the diversity and numbers of species in both the perciforms and scorpaeniforms may continue to deter their resolution.

Within the suborder Scorpaenoidei, Ishida (1994) considered the Sebastidae to contain eight genera. Some members of the Sebastidae have derived states of 9 of the 95 characters that he used to examine the Scorpaenoidei (Tables 3, 4). All members of the family have derived states of two characters relative to their states in lower percoids, the presumed outgroup of scorpaenoids: the shortening of al fibers with the lengthening of the associated maxillary tendon (character 3), and an increase in the number of vertebrae from 24 (character 70) (Table 3).

Within the Sebastidae, the genera Sebastes, Sebastiscus, and Hozukius share identical character states, resulting in an unresolved trichotomy, which can be given the subfamilial name Sebastinae. This subfamily has the derived states of four characters and shares the derived states of two of these characters with all of the other genera in Sebastidae. However, they do not possess uniquely derived states of any characters. They are united in possessing swimbladders, which is a character reversal (in that the rest of the family lacks them, which is the presumed derived state). If Hozukius is found to be viviparous like Sebastes and Sebastiscus, then viviparity would be another derived character to support this subfamily. If Hozukius is found to be oviparous, it may be considered the primitive sister genus of Sebastes and Sebastiscus within the subfamily.

What Morphological Characters Can Be Used for Cladistic Analysis of Sebastes?

To be used in a cladistic analysis, morphological characters must be independent of each other and nonadaptive. However, many of the characters used to identify Sebastes and used in early attempts to establish relationships among the species are correlated with each other and are adaptive for either pelagic or demersal existence (Table 5).

There is a basic dichotomy in Sebastes ecology: species either live in the water column or on the bottom. This is not an absolute; there is a cline between these two basic conditions. Bottom-dwelling species have the lower jaw protruding, eyes close together with a concave interorbital area, heavy armature, thickened pectoral rays, short gill rakers, scaled jaws, and a curved skull base, while pelagic species have opposite states for all of these characters. Convergence toward these character states is expected in species occupying these niches regardless of their ancestry.

Another factor to consider when interpreting characters is that meristic characters tend to show a latitudinal cline: the closer the species lives to the pole, the higher the value (Jordan's Rule: see Lindsey, 1988). This basic pattern is seen in medial fin ray and vertebral counts among species of rockfishes (Table 5). For example, Sebastiscus, which lives farther south than most other taxa, has low counts for all these characters and Emmelas, which lives in the northern part of the range, has higher counts. The species of Sebastomus, which live primarily in the southern part of the range of Sebastes in the northeast Pacific, including the Gulf of California, and the Southern Hemisphere, have lower medial fin ray counts than most other Sebastes. The Atlantic species of Sebastes have the highest vertebral counts in the genus, and presumably they originated from species that traversed the Arctic Ocean from the Pacific.

Can Early Life History Information Be Applied to the Problem?

Scorpaenoids display a variety of reproductive patterns, but the use of information on reproduction in systematic studies is limited, since the patterns are unknown in several genera (Washington et al., 1984b). While most scorpaenoids share ovoviviparity with most other teleosts, Sebastes, Sebastiscus, and possibly Helicolenus share the unique pattern of internal fertilization, intraovarian embryological development with maternal contribution to nutrition, and production of many ([~10.sup.5]) newly hatched, undeveloped larvae. The reproductive pattern of Hozukius is unknown, although the anal anatomy is that of a live-bearer (Ida (4)). As opposed to the widespread pattern of producing individual planktonic eggs, several scorpaenid genera (Sebastolobus, Pterois, Scorpaena) extrude eggs in a floating bilobed gelatinous egg mass. A single oil globule is present in eggs of Pterois and Sebastolobus, while none is present in the eggs of Scorpaena or Dendrochirus.

The larvae of many scorpaeniforms, including the scorpaenoids, have parietal and other head spines, which are not seen in larvae of other fishes. In Sebastolobus and Scorpaenodes the parietal ridge is enlarged and ends in a bifid spine, with the posterior part (the nuchal spine) being predominant, whereas in Sebastes, Sebastiscus (Okiyama, 1988), and Helicolenus the parietal spine is larger and slightly separate from the nuchal spine (Moser et al., 1977; Moser, 1996). There is a tendency for early development of pectoral fins in scorpaeniforms, and in some groups the pectoral fins are quite pronounced in the larvae. Pronounced pectoral fins are seen in Sebastolobus, Scorpaenodes, Ectreposebastes and some species of Sebastes (e.g. S. levis). Larvae of Helicolenus develop a spongy mass of tissue anteriorly along the dorsal midline. Once the larvae of more scorpaenoids become known, such characters will prove invaluable in systematic studies of the group.

Within the genus Sebastes, the preflexion-extrusion larvae of most species are known (Matarese et al., 1989; Moser, 1996). These can be obtained fairly easily, and their identity known, by collecting pregnant females and extruding the larvae. Aside from some shared characteristics among preflexion larvae of species of Sebastomus, the appearance of these larvae seems to offer little taxonomic or systematic promise. There is considerable overlap in characters among species in these larvae and considerable within-species variation.

Later-stage larvae of Sebastes are distinctive from larvae of other fishes, but many share characters with each other that prevent identifying them to species in all but a few cases. Larval characters that may be useful in taxonomy and systematics include body shape, pigment patterns, and the development and subsequent reduction or loss of head spines (Kendall, 1991). As with early larvae, the later larvae of Sebastomus sham some characters that distinguish them from other Sebastes larvae, but within the subgenus, the larvae of many species are quite similar. It appears that larvae of Pteropodus may also share some characters, but the larvae of several species are still unknown. A few other species have distinctive larvae (e.g. Sebastes paucispinis, S. jordani, S. melanostomus, and S. levis), but their morphology does not seem to be related to their subgeneric placement. There are some intriguing similarities among the larvae of some species of Sebastes that may indicate relationships not otherwise suggested (e.g. the larvae of S. jordani, S. alutus, and S. polyspinis from the Pacific all look quite similar to the larvae of the Atlantic species of Sebastes).

As juveniles, Sebastes are morphologically quite different from the larvae and the adults. Some species reside in the water column, some associate with flotsam or school, and some take up a demersal existence (Moser and Boehlert, 1991). The observed pigment patterns of juveniles may be adaptive for these various modes of existence, so caution is advised when using them in systematic analysis.

Clearly the early life history of scorpaenoids in general and Sebastes in particular offers a large number of characters to apply to systematic studies. However, until the reproductive patterns and early life history series of more species are described, this potential cannot be realized.

Conclusions

After a period of discovering the incredible diversity of Sebastes in the middle 1800's, the history of the systematics of the genus was marked by bitter debate in the late 1800's over relationships among the species. With few exceptions (e.g. S. carnatus-S, chrysomelas-S, atrovirens, and S. ciliatus) the species of Sebastes are now well-defined, and few new ones are being described (Kim and Lee, 1994; Eitner et al., 1999). However, little recent work has been done on the genus as a unit, and its systematics remain in a confused state, with over 100 species, and very little obvious structure within the genus. What other fish genus contains so many species, but with such little indication of structure within it? Beyond the fairly cohesive group Sebastomus and a few species pairs (e.g. Sebastes fasciatus/S. mentella, S. babcocki/S, rubrivinctus, S. serranoides/S. flavidus), practically any two species look as similar to each other as to any other species.

In order to progress in our understanding of the relationships within this interesting, and ecologically and economically important group of fishes, traditional gross morphological as well as novel characters, such as those avail able through genetics, need to be evaluated more rigorously. With the incredible size of the genus and its wide distributional range, this will be an enormous task.

Appendix I

Original Definitions of Sebastes, Hozukius, and Sebastiscus, and the Subgenera of Sebastes

Sebastes Cuvier (1829), type species: S. norvegicus: "Have all the characters of the Scorpaena except that they lack cutaneous filaments, and that their head, less covered with bristles, is scaly.

"There is a large species of them in the North Sea, called `marulke,' and in some places `carp' (Sebastes norvegicus, Nob., Perca marina, Pennt., Perca norvegica, Mull.), Bonnat. Encycl. Meth., plate on ichthyology, fig. 210. It is red, and is often more than two feet long. It is dried to make food provisions. Its dorsal spines are used as needles by Eskimos.

"The Mediterranean has a species very similar, but its dorsal rays are less numerous (Sebastes imperialis, Nob., Scorpaena dactyloptera, Laroche, Annales Mus., 13, plate 22, fig. 9). Its palate is black; it lacks a swim bladder, although the preceding species has one." (2)

Sebastodes Gill (1861), type species: S. paucispinis: "A very different facies from Sebastes, and is readily distinguished by the longer body, the very protuberant lower jaw, which has a symphyseal swelling beneath, the very minute scales, the form and armature of the head, the deep emargination of the dorsal fin and the emarginated caudal."

Sebastichthys Gill (1862b), type species: S. nigrocinctus: "Species referred to the genus Sebastes, which has eleven to twelve (XI.+1.-XII.+1.) spines in its first dorsal fin, palatine teeth, and the physiognomy of Sebastes (norvegicus)."

Sebastomus Gill (1864), type species: S. rosaceus: "Distinguished by the texture of the bones of the skull, armed orbital ridges, prefrontals &c."

Sebastosomus Gill (1864), type species: S. melanops: "... the genus Sebastichthys includes at least three genera. The Sebastichthys nigrocinctus is somewhat related to Scorpaena, and distinguished by elevated, serrated coronal crests. The other California species represented by the Sebastes melanops, seen by me, differ so much that they may be separated and combined under the genus Sebastosomus ..."

Acutomentum Eigenmann and Beeson (1893), type species: S. ovalis: Taken from their key: "Parietals meeting above the supra-occipital. Lower jaw much projecting; head broad, the skull usually convex; cranial ridges, when present, low; gill rakers very long and slender; scales usually smooth, few if any accessory scales. Parietal ridges ending in spines; preocular, supraocular and tympanic spines well developed. Peritoneum black. Postocular spine present. Second anal spine usually stronger and longer than third. Symphyseal knob strong, projecting forward. Dorsal low. (Peritoneum black, mandibles and maxillary scaled)."

Primospina Eigenmann and Beeson (1893), type species: S. mystinus: Taken from their key: Parietals meeting above the supra-occipital. Lower jaw much projecting; head broad, the skull usually convex; cranial ridges, when present, low; gill rakers very long and slender; scales usually smooth, few if any accessory scales. Postocular spine not developed. Parietal ridges not ending in spines. Preocular spines well developed. Supraocular and tympanic spines sometimes present. Interorbital wide, convex. Peritoneum black. Approximated edges of sub-opercle, and interopercle frequently ending in spines.

Pteropodus Eigenmann and Beeson (1893), type species: S. maliger: Taken from their key: Parietals separated by the supra-occipital. Cranium with many ridges, all ending in spines. Postocular spines wanting. Coronal spines none.

Auctospina Eigenmann and Beeson (1893), type species: S. auriculatus: Taken from their key: Parietals separated by the supra-occipital. Cranium with many ridges, all ending in spines. Postocular spines wanting. Coronal spines present.

Eosebastes Jordan and Evermann (1896), type species: S. aurora: No description of the subgenus offered; see description of type species in Jordan and Evermann (1898).

Hispaniscus Jordan and Evermann (1896), type species: S. rubrivinctus: No description of the subgenus offered; see description of type species in Jordan and Evermann (1898).

Rosicola Jordan and Evermann (1896), type species: S. pinniger: No description of the subgenus offered; see description of type species in Jordan and Evermann (1898).

Emmelas Jordan and Evermann (1898), type species: S. glaucus: No description of the subgenus offered; see description of type species in Jordan and Evermann (1898).

Zalopyr Jordan and Evermann (1898), type species: S. aleutianus: No description of the subgenus offered; see description of type species in Jordan and Evermann (1898).

Sebastiscus Jordan and Starks (1904), type species: Sebastiscus marmoratus: "This genus is based on species having the external appearance of Sebastodes and much resemblance to the subgeneric group called Pteropodus, but having 12 spines in the dorsal fin and the vertebrae 10+14=24, agreeing in these regards with Scorpaena. From Helicolenus, Sebastiscus differs, solely, perhaps, in the presence of a well-developed airbladder. Peritoneum pale."

Sebastocles Jordan and Hubbs (1925), type species: S. elegans (=S. hubbsi): "Dorsal spines low, normally fourteen; interorbital deeply concave, size small."

Sebastocarus Jordan and Evermann (1927), type species: S. serriceps: "This genus is closely allied to Sebastichthys from which it differs in the more compressed body and especially in the strict and high ridges on the head, all of them being free from serrations or accessory tubercles."

Sebastopyr Jordan and Evermann (1927), type species: S. ruberrimus: "This genus is allied to Sebastomus, but of much coarser build, the cranial spines in the adult being rough with blunt spinules."

Hozukius Matsubara (1934), type species: H. emblemarius: "External: Body rather robust and somewhat compressed. Maxillary, and mandible covered with small ctenoid scales but not on branchiostegals. Teeth in villiform bands on vomer and palatines. Head much spinous; nasal, preocular, supraocular, postocular, tympanic, coronal, parietal, and nuchal spines present; upper and posterior margins of orbit provided with many accessory spines; parietal ridge not conspicuous. Orbital margins of preorbital and second suborbital bones armed with a sharp spine which is directed backward; first suborbital margined with several small spines; lower border of preorbital with two very strong, sharp bifid spines. Lower three preopercular spines much larger than upper two; subopercle and interopercle each with a small, sharp spine at their approximation. Symphyseal knob very large. Dorsal with 12 spines and about 12 rays; anal with 3 spines and about 6 rays. Auxiliary flap absent.

"Internal: Skull thick and heavy. Base of skull (paraspheniod) moderately curved. Frontal, parietal, pterotic and supratemporal much cavernous resembling a bee-hive in general appearance. Parietals entirely meeting; upper margin of orbit and frontal ridges somewhat raised. Mesethmoid processes directed upward and forward. Nodule on the front of prefrontal for the articulation with palatine well developed. Ventral process of basisphenoid rudimentary, not reaching to paraspheniod; basisphenoid with a small pore between it and basis crania. Myodome well developed, with a rather large posterior opening. Second suborbital bone very broad and long entirely touching to the upper ridge of preopercle; third and fourth suborbital bones present. Gill-rakers long and stout, the longest one much longer than the longest gilllamella. Vertebrae 9 + 16 + hypural = 26; parapophysis developed from the sixth vertebra, its processes widely diverging, directed outward and downward. Air bladder absent. Peritoneum jet black."

Hatumeus Matsubara (1943), type species: S. owstoni: "Cranium thin and papery; interorbital space and occiput flat, the former about 3.3 times in base of the cranium; preocular, postocular, tympanic and parietal spines present, but small and weak; nuchal spines sometimes present; supraocular and coronal spines absent; cranial ridges absent except for the parietal ridges which are low but never scaled over; parietals separated; mesethmoid processes never directed upward; base of cranium markedly curved; ventral process of basisphenoid feeble, never attached to the parasphenoid; nasal spines low, but strong; preorbital lobes without spines; second suborbital bone much wider than long. Vertebrae usually 30 including the hypural. Peritoneum jet black. Gill-rakers long and slender. Dorsal mostly XIV, 14; anal usually III, 9 to 10. Lower jaw strongly produced beyond the upper when the mouth is closed; symphyseal knob evident; jaws thickly covered by small scales."

Mebarus Matsubara (1943), type species: S. inermis: "Cranium rather thin; interorbital space and occiput plain or very slightly convex, the former about 3.0 times in base of the cranium; preocular spines always present; supraocular and parietal spines small or absent; postocular, tympanic and nuchal spines entirely absent; cranial ridges absent except for the parietal which is very low and sometimes scaled over; parietals widely separated; mesethmoid processes depressed or slightly directed upward; base of cranium strongly curved; ventral process of basisphenoid feeble; nasal spines low, but strong. Vertebrae usually 26 or 27 including the hypural. Peritoneum black or pale. Gill-rakers long and slender. Dorsal spines 13. Lower jaw projecting far beyond the upper, provided with a prominent symphyseal knob; maxillary at least scaly; preorbital lobes with or without spines.

"The present new subgenus is closely related to the subgenus Rosicola, but differs from it in having weak parietal ridges and flatfish interorbital space and lacking the postocular and tympanic spines. The subgenus is also separable from Sebastosomus in having usually the strong preocular spine, flatfish interorbital space and parietals widely separated."

Murasoius Matsubara (1943), type species: S. pachycephalus: "Cranium very thick and heavy; interorbital space deeply concave, about 3 1/2 in base of cranium; cranial spines broad and stout, the pointed tips directed backward; preocular, postocular, tympanic and parietal spines present; supraocular, coronal and nuchal spines absent; supraocular ridges exceedingly high and stout, forming a narrow flat area between them; frontal ridges barely evident, can not be seen without elimination of the skin; parietal ridges broad and high, but a little lower than the supraocular ridges; nasal spines stout, directed upward and slightly backward; mesethmoid processes strongly compressed, directed forward and upward; parietal bones separated; base of cranium straight; ventral process of basisphenoid entirely meeting the parasphenoid. Vertebrae 26 including hypural. Peritoneum white. Air-bladder entirely free from the layer of connective tissue lying outside the peritoneum, the bladder being easily detached from the coelom; the extrinsic muscle band of the bladder gives rise posteriorly to a single short ligament; the anterior part of the muscle band not pierced through by a ligament. Gill-rakers very short and blunt. Dorsal spines 13. Jaws, snout and branchiostegals mostly naked; upper jaw projecting beyond the lower when mouth is closed; symphyseal knob at the tip of lower jaw absent or barely evident."

Neohispaniscus Matsubara (1943), type species: S. schlegelii: "Skull thick and heavy; interorbital space flat or slightly convex, its width about 2.6 to 3.3 times in base of the cranium; cranial spines well developed, directed straightly backward; preocular, postocular, tympanic and parietal spines always present supraocular, corneal and nuchal spines absent; parietal ridges high and naked; frontal ridges low but distinct, the space between them shallowly concave; supraocular edges low or depressed, as high as or lower than the frontal ridges; mesethmoid processes slightly elevated upward; base of cranium some-what curved; parietals separated or partly meeting; ventral process of the basisphenoid well developed, entirely meeting the parasphenoid. Vertebrae 26 including hypural. Peritoneum usually white. Gill-rakers, comparatively long and slender with pointed tips. Lower jaw generally projecting beyond the upper when the mouth is closed, with a rather small symphyseal knob. Jaws usually naked.

"The present subgenus is closely related to Hispaniscus, but differs from it in having a much longer and wider inoterorbital space, elevated mesethmoid processes and shallowly depressed area between the frontal ridges."

Takenokius Matsubara (1943), type species: S. oblongus: "Cranium thick and heavy; interorbital space and occiput flat or slightly convex, the former about 3.7 times in base of cranium; postocular, tympanic and parietal spines present, directed backward and downward; preocular supraocular, coronal and nuchal spines entirely absent; parietal ridges low but broad; supraocular edges depressed, never higher than the frontal ridges; frontal ridges low, the space between them flattish, never deeply concave; mesethmoid processes directed forward and upward; parietals meeting or narrowly separated; the patch of vomerine teeth triangular; base of cranium straight; ventral process of basisphenoid meeting the parasphenoid. Vertebrae 26 including hypural. Peritoneum pale. Gill-rakers very short, tubercular. Jaws equal in length and entirely scaleless; symphyseal knob inconspicuous."

Allosebastes Hubbs (1951), type species: S. sinensis: "... reduction of the anal soft-rays to 5.... smooth, mostly cycloid scales; the unswollen lower pectoral rays; the excessively long anal spines (the second extends well beyond the longest soft-ray); the closely clumped subparallel upper 3 preopercular spines (the lower 2 are well separated and divergent); the protuberant posterior end of the mandible, almost resembling a flat spine; and, especially, the very firm well-exposed suborbital stay (second suborbital), with the bone widened about the unusually large pore, which has a somewhat raised and roughened rim."

Appendix II

Summary of Nomenclature
of Sebastes Listed Alphabetically
by Subgenus

Original species (1) Authors Date

 (Fitch) 1964
 Lea and Fitch 1979
 (Westrheim and Tsuyuki) 1967
 Barsukov 1988
 (Taranetz and Moiseev) 1933
 Kim and Lee 1994
 Eitner et al. 1999

alutum (Gilbert) 1890
brevispine (Bean) 1884
clavilatum Starks 1911
eigenmanni Cramer 1896
entomelas (Jordan and Gilbert) 1880
 (Cramer) 1895
 (Eigenmann and Beeson) 1893
ovale (Ayres) 1863
rufum (Eigenmann and Eigenmann) 1890
 (Jordan and Starks) 1904
 (Jordan and Starks) 1904
 (Jordan and Snyder) 1900
 (Wakiya) 1917
paucispinosus Matsubara 1943

 Chen 1975
 Chen 1975
 (Beebe and Tee-Van) 1938
 (Gilbert) 1890
 Quast 1971
 Lea and Fitch 1972
 (Gilbert) 1890
 (Starks) 1911
prorigerum (Jordan and Gilbert) 1880
 (Gilbert) 1890
 (Gilbert) 1897
 (Gilbert) 1915
 (Gilbert) 1890

dallii (Eigenmann and Beeson) 1894
auriculata Girard 1854

 Hilgendorf 1880

 (Gilbert) 1890
 (Jordan) 1897
deani Starks 1911
introniger (Gilbert) 1890
 (Eigenmann and Eigenmann) 1890
rupestris (Gilbert) 1890

 (Jordan and Thompson) 1914

 Ayres 1859
 (Eigenmann and Eigenmann) 1889
 (Jordan and Gilbert) 1880
atrorubens (Gilbert) 1898
 (Jordan and Gilbert) 1880
 (Jordan and Hubbs) 1925
 (Matsubara) 1934
 Cuvier and Valenciennes 1829
 Gunther 1878
taczanowskii (Steindachner) 1880
 Barsukov 1972
guentheri (Jordan and Starks) 1904
paradoxus Matsubara 1943
tokionis (Jordan and Starks) 1904

 Matsubara 1943
 Temminck and Schlegel 1843
 (Schmidt) 1931
 Matsubara 1943

 Hilgendorf 1880
 Steindachner and Doderlein 1884

ijimae (Jordan and Metz) 1913
zonatus Chen and Barsukov 1976

 (Jordan and Gilbert) 1880
 Richardson 1844
 (Jordan and Gilbert) 1881
gilberti Cramer 1896
 (Jordan and Gilbert) 1880
 Ayres 1854
 (Jordan and Gilbert) 1880
vexillaris (Jordan and Gilbert) 1880
 Hilgendorf 1880
 Hilgendorf 1880
 (Matsubara) 1937
 (Matsubara) 1934

alexandri (Evermann and Goldsborough) 1907
 (Thompson) 1915
 (Jordan and Gilbert) 1880
 (Gill) 1864

 (Travin) 1951
 Storer 1854
norvegica (Ascanius) 1772
 Kroyer 1845

 Ayres 1859

albo-fasciatus (Lacepede) 1802
 (Cuvier and Valenciennes) 1829
tertius (Barsukov and Chen) 1978

 (Jordan and Gilbert) 1880

 (Eigenmann and Eigenmann) 1890
 (Gilbert) 1896
 Ayres 1854
 (Jordan and Starks) 1904
 Hilgendorf 1880

 Chen 1975
 Chen 1971
 Chen 1971
 Chen 1971
 Chen 1971
 Chen 1971
 Ayres 1859
 Chen 1971
ayresii (Gilbert and Cramer) 1897
 (Jordan and Gilbert) 1880
 (Jordan and Gilbert) 1880
 (Eigenmann and Eigenmann) 1890
gillii (Eigenmann) 1891
rhodochloris (Jordan and Gilbert) 1880
 Girard 1854
 (Jordan and Gilbert) 1882
 Valenciennes 1833
 (Gmelin) 1789
chamaco (Evermann and Radcliffe) 1917

 (Tilesius) 1813
 (Ayres) 1862
 Girard 1856
 (Jordan and Gilbert) 1881
 (Eigenmann and Eigenmann) 1890

 (Jordan and Gilbert) 1883

 Gunther 1877

 Barsukov 1970
 (Jordan and Evermann) 1898
kawaradai (Matsubara) 1934
melanostictus (Matsubara) 1934
swifti (Evermann and Goldsborough) 1906
 Hilgendorf 1880

 Present
Original species (1) Authors species (2)

 (Fitch) phillipsi
 Lea and Fitch melanosema
 (Westrheim and Tsuyuki) reedi
 Barsukov cheni
 (Taranetz and Moiseev) polyspinis
 Kim and Lee koreanus
 Eitner et al. moseri

alutum (Gilbert) alutus
brevispine (Bean) brevispinis
clavilatum Starks (?)
eigenmanni Cramer (?)
entomelas (Jordan and Gilbert) entomelas
 (Cramer) hopkinsi
 (Eigenmann and Beeson) macdonaldi
ovale (Ayres) ovalis
rufum (Eigenmann and Eigenmann) rufus
 (Jordan and Starks) flammeus
 (Jordan and Starks) iracundus
 (Jordan and Snyder) scythropus
 (Wakiya) baramenuke
paucispinosus Matsubara (alutus)

 Chen varispinis
 Chen peduncularis
 (Beebe and Tee-Van) cortezi
 (Gilbert) sinensis
 Quast variegatus
 Lea and Fitch rufinanus
 (Gilbert) diploproa
 (Starks) emphaeus
prorigerum (Jordan and Gilbert) proriger
 (Gilbert) saxicola
 (Gilbert) semicinctus
 (Gilbert) wilsoni
 (Gilbert) zacentrus

dallii (Eigenmann and Beeson) dalli
auriculata Girard auriculatus

 Hilgendorf glaucus

 (Gilbert) aurora
 (Jordan) crameri
deani Starks (?)
introniger (Gilbert) (melanostomus)
 (Eigenmann and Eigenmann) melanostomus
rupestris (Gilbert) (melanostomus)

 (Jordan and Thompson) owstoni

 Ayres elongatus
 (Eigenmann and Eigenmann) levis
 (Jordan and Gilbert) rubivinctus
atrorubens (Gilbert) (atrovirens)
 (Jordan and Gilbert) atrovirens
 (Jordan and Hubbs) thompsoni
 (Matsubara) wakiyai
 Cuvier and Valenciennes inermis
 Gunther joyned
taczanowskii (Steindachner) taczanowski
 Barsukov minor
guentheri (Jordan and Starks) (inermis)
paradoxus Matsubara (wakiyai)
tokionis (Jordan and Starks) (inermis)

 Matsubara * nudus
 Temminck and Schlegel pachycephalus
 (Schmidt) * nigricans
 Matsubara * chalcogrammus

 Hilgendorf schlegeli
 Steindachner and Doderlein vulpes

ijimae (Jordan and Metz) (vulpes)
zonatus Chen and Barsukov (vulpes)

 (Jordan and Gilbert) carnatus
 Richardson caurinus
 (Jordan and Gilbert) chrysomelas
gilberti Cramer (dalli)
 (Jordan and Gilbert) maliger
 Ayres nebulosus
 (Jordan and Gilbert) rastrelliger
vexillaris (Jordan and Gilbert) (caurinus)
 Hilgendorf nivosus
 Hilgendorf trivittatus
 (Matsubara) hubbsi
 (Matsubara) longispinis

alexandri (Evermann and Goldsborough) (auriculatus)
 (Thompson) babcocki
 (Jordan and Gilbert) miniatus
 (Gill) pinniger

 (Travin) mentella
 Storer fasciatus
norvegica (Ascanius) norvegicus
 Kroyer viviparus

 Ayres nigrocinctus

albo-fasciatus (Lacepede) albofasciatus
 (Cuvier and Valenciennes) marmoratus
tertius (Barsukov and Chen) (marmoratus)

 (Jordan and Gilbert) serriceps

 (Eigenmann and Eigenmann) goodei
 (Gilbert) jordani
 Ayres paucispinis
 (Jordan and Starks) itinus
 Hilgendorf steindachneri

 Chen spinorbis
 Chen exsul
 Chen simulator
 Chen rosenblatti
 Chen ensifer
 Chen notius
 Ayres helvomaculatus
 Chen lentiginosus
ayresii (Gilbert and Cramer) (rosaceus)
 (Jordan and Gilbert) chlorostictus
 (Jordan and Gilbert) constellatus
 (Eigenmann and Eigenmann) eos
gillii (Eigenmann) gilli
rhodochloris (Jordan and Gilbert) (helvomaculatus)
 Girard rosaceus
 (Jordan and Gilbert) umbrosus
 Valenciennes oculatus
 (Gmelin) capensis
chamaco (Evermann and Radcliffe) (capensis)

 (Tilesius) ciliatus
 (Ayres) flavidus
 Girard melanops
 (Jordan and Gilbert) mystinus
 (Eigenmann and Eigenmann) serranoides

 (Jordan and Gilbert) ruberrimus

 Gunther oblongus

 Barsukov borealis
 (Jordan and Evermann) aleutianus
kawaradai (Matsubara) (aleutianus)
melanostictus (Matsubara) (aleutianus)
swifti (Evermann and Goldsborough) (aleutianus)
 Hilgendorf matsubarae

Original species (1) Authors Common name

 (Fitch) chameleon
 rockfish
 Lea and Fitch semaphore
 rockfish
 (Westrheim and Tsuyuki) yellowmouth
 rockfish
 Barsukov
 (Taranetz and Moiseev) northern
 rockfish
 Kim and Lee hwanghee-bolnak
 Eitner et al. whitespeckled
 rockfish

alutum (Gilbert) Pacific ocean
 perch
brevispine (Bean) silvergray
 rockfish
clavilatum Starks
eigenmanni Cramer
entomelas (Jordan and Gilbert) widow rockfish
 (Cramer) squarespot
 rockfish
 (Eigenmann and Beeson) Mexican rockfish
ovale (Ayres) speckled
 rockfish
rufum (Eigenmann and Eigenmann) bank rockfish
 (Jordan and Starks) sankou-menuke
 (Jordan and Starks) oo-saga
 (Jordan and Snyder) ukeguchi-mebaru
 (Wakiya) bara-menuke
paucispinosus Matsubara

 Chen
 Chen
 (Beebe and Tee-Van)
 (Gilbert)
 Quast harlequin
 rockfish
 Lea and Fitch dwarf-red
 rockfish
 (Gilbert) splitnose
 rockfish
 (Starks) Puget Sound
 rockfish
prorigerum (Jordan and Gilbert) redstripe
 rockfish
 (Gilbert) stripetail
 rockfish
 (Gilbert) halfbanded
 rockfish
 (Gilbert) pygmy rockfish
 (Gilbert) sharpchin
 rockfish

dallii (Eigenmann and Beeson) calico rockfish
auriculata Girard brown rockfish

 Hilgendorf kuro-menuke

 (Gilbert) aurora rockfish
 (Jordan) darkblotched
 rockfish
deani Starks
introniger (Gilbert)
 (Eigenmann and Eigenmann) blackgill
 rockfish
rupestris (Gilbert)

 (Jordan and Thompson) hatsume

 Ayres greenstriped
 rockfish
 (Eigenmann and Eigenmann) cowcod
 (Jordan and Gilbert) flag rockfish
atrorubens (Gilbert)
 (Jordan and Gilbert) kelp rockfish
 (Jordan and Hubbs) usu-mebaru
 (Matsubara) gaya-modoki
 Cuvier and Valenciennes mebaru
 Gunther togotto-mebaru
taczanowskii (Steindachner) ezo-mebaru
 Barsukov aka-gaya
guentheri (Jordan and Starks)
paradoxus Matsubara
tokionis (Jordan and Starks)

 Matsubara
 Temminck and Schlegel mura-soi
 (Schmidt)
 Matsubara

 Hilgendorf kuro-soi
 Steindachner and Doderlein kitsune-mebaru

ijimae (Jordan and Metz)
zonatus Chen and Barsukov

 (Jordan and Gilbert) gopher rockfish
 Richardson copper rockfish
 (Jordan and Gilbert) black-and-yellow
 rockfish
gilberti Cramer
 (Jordan and Gilbert) quillback
 rockfish
 Ayres China rockfish
 (Jordan and Gilbert) grass rockfish
vexillaris (Jordan and Gilbert)
 Hilgendorf goma-soi
 Hilgendorf shima-zoi
 (Matsubara) yoroi-mebaru
 (Matsubara) kourai-yoroi
 mebaru

alexandri (Evermann and Goldsborough)
 (Thompson) redbanded
 rockfish
 (Jordan and Gilbert) vermilion
 rockfish
 (Gill) canary rockfish

 (Travin) deepwater
 redfish
 Storer Acadian redfish
norvegica (Ascanius) golden redfish
 Kroyer Norway haddock

 Ayres tiger rockfish

albo-fasciatus (Lacepede) ayame-kasago
 (Cuvier and Valenciennes) kasago
tertius (Barsukov and Chen)

 (Jordan and Gilbert) treefish

 (Eigenmann and Eigenmann) chilipepper
 (Gilbert) shortbelly
 rockfish
 Ayres bocaccio
 (Jordan and Starks) yanagi-mebaru
 Hilgendorf yanagi-no-mai

 Chen
 Chen
 Chen pinkrose
 rockfish
 Chen greenblotched
 rockfish
 Chen swordspine
 Chen rockfish
 Ayres rosethorn
 rockfish
 Chen freckled
 rockfish
ayresii (Gilbert and Cramer)
 (Jordan and Gilbert) greenspotted
 rockfish
 (Jordan and Gilbert) starry rockfish
 (Eigenmann and Eigenmann) pink rockfish
gillii (Eigenmann) bronzespotted
 rockfish
rhodochloris (Jordan and Gilbert)
 Girard rosy rockfish
 (Jordan and Gilbert) honeycomb
 rockfish
 Valenciennes
 (Gmelin)
chamaco (Evermann and Radcliffe)

 (Tilesius) dusky rockfish
 (Ayres) yellowtail
 rockfish
 Girard black rockfish
 (Jordan and Gilbert) blue rockfish
 (Eigenmann and Eigenmann) olive rockfish

 (Jordan and Gilbert) yelloweye
 rockfish

 Gunther takenoko-mebaru

 Barsukov shortraker
 rockfish
 (Jordan and Evermann) rougheye
 rockfish
kawaradai (Matsubara)
melanostictus (Matsubara)
swifti (Evermann and Goldsborough)
 Hilgendorf akou-dai
 Majorgeneric or
 subgeneric
 assignments
 Eigenmann and
Original species (1) Authors Beeson, 1894

 (Fitch)
 Lea and Fitch
 (Westrheim and Tsuyuki)
 Barsukov
 (Taranetz and Moiseev)
 Kim and Lee
 Eitner et al.

alutum (Gilbert) Acutomentum
brevispine (Bean) Pteropodus
clavilatum Starks
eigenmanni Cramer
entomelas (Jordan and Gilbert) Primospina
 (Cramer)
 (Eigenmann and Beeson) Acutomentum
ovale (Ayres) Acutomentum
rufum (Eigenmann and Eigenmann) Sebastomus
 (Jordan and Starks)
 (Jordan and Starks)
 (Jordan and Snyder)
 (Wakiya)
paucispinosus Matsubara

 Chen
 Chen
 (Beebe and Tee-Van)
 (Gilbert) Pteropodus
 Quast
 Lea and Fitch
 (Gilbert) Sebastichthys
 (Starks)
prorigerum (Jordan and Gilbert) Pteropodus
 (Gilbert) Pteropodus
 (Gilbert)
 (Gilbert)
 (Gilbert) Pteropodus

dallii (Eigenmann and Beeson) Pteropodus
auriculata Girard Auctospina

 Hilgendorf

 (Gilbert) Auctospina
 (Jordan)
deani Starks
introniger (Gilbert) Acutomentum
 (Eigenmann and Eigenmann) Acutomentum
rupestris (Gilbert) Sebastomus

 (Jordan and Thompson)

 Ayres
 (Eigenmann and Eigenmann) Sebastomus
 (Jordan and Gilbert) Sebastichthys
atrorubens (Gilbert)
 (Jordan and Gilbert) Pteropodus
 (Jordan and Hubbs)
 (Matsubara)
 Cuvier and Valenciennes
 Gunther
taczanowskii (Steindachner)
 Barsukov
guentheri (Jordan and Starks)
paradoxus Matsubara
tokionis (Jordan and Starks)

 Matsubara
 Temminck and Schlegel
 (Schmidt)
 Matsubara

 Hilgendorf
 Steindachner and Doderlein

ijimae (Jordan and Metz)
zonatus Chen and Barsukov

 (Jordan and Gilbert) Pteropodus
 Richardson Pteropodus
 (Jordan and Gilbert) Pteropodus
gilberti Cramer
 (Jordan and Gilbert) Pteropodus
 Ayres Pteropodus
 (Jordan and Gilbert) Pteropodus
vexillaris (Jordan and Gilbert) Pteropodus
 Hilgendorf
 Hilgendorf
 (Matsubara)
 (Matsubara)

alexandri (Evermann and Goldsborough)
 (Thompson)
 (Jordan and Gilbert) Sebastomus
 (Gill) Sebastomus

 (Travin)
 Storer
norvegica (Ascanius)
 Kroyer

 Ayres Sebastichthys

albo-fasciatus (Lacepede)
 (Cuvier and Valenciennes)
tertius (Barsukov and Chen)

 (Jordan and Gilbert) Sebastichthys

 (Eigenmann and Eigenmann) Sebastodes
 (Gilbert)
 Ayres Sebastodes
 (Jordan and Starks)
 Hilgendorf

 Chen
 Chen
 Chen
 Chen
 Chen
 Chen
 Ayres Sebastomus
 Chen
ayresii (Gilbert and Cramer)
 (Jordan and Gilbert) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Eigenmann and Eigenmann) Sebastomus
gillii (Eigenmann) Sebastomus
rhodochloris (Jordan and Gilbert) Sebastomus
 Girard Sebastomus
 (Jordan and Gilbert) Sebastomus
 Valenciennes
 (Gmelin)
chamaco (Evermann and Radcliffe)

 (Tilesius) Sebastosomus
 (Ayres) Sebastosomus
 Girard Sebastosomus
 (Jordan and Gilbert) Primospina
 (Eigenmann and Eigenmann) Sebastosomus

 (Jordan and Gilbert)

 Gunther

 Barsukov
 (Jordan and Evermann)
kawaradai (Matsubara)
melanostictus (Matsubara)
swifti (Evermann and Goldsborough)
 Hilgendorf
 Majorgeneric or
 subgeneric
 assignments

 Jordan and
Original species (1) Authors Evermann, 1898

 (Fitch)
 Lea and Fitch
 (Westrheim and Tsuyuki)
 Barsukov
 (Taranetz and Moiseev)
 Kim and Lee
 Eitner et al.

alutum (Gilbert) Acutomentum
brevispine (Bean) Acutomentum
clavilatum Starks
eigenmanni Cramer Acutomentum
entomelas (Jordan and Gilbert) Acutomentum
 (Cramer) Acutomentum
 (Eigenmann and Beeson) Acutomentum
ovale (Ayres) Acutomentum
rufum (Eigenmann and Eigenmann) Acutomentum
 (Jordan and Starks)
 (Jordan and Starks)
 (Jordan and Snyder)
 (Wakiya)
paucispinosus Matsubara

 Chen
 Chen
 (Beebe and Tee-Van)
 (Gilbert) Hispaniscus
 Quast
 Lea and Fitch
 (Gilbert) Eosebastes
 (Starks)
prorigerum (Jordan and Gilbert) Acutomentum
 (Gilbert) Eosebastes
 (Gilbert) Eosebastes
 (Gilbert)
 (Gilbert) Hispaniscus

dallii (Eigenmann and Beeson)
auriculata Girard Auctospina

 Hilgendorf Emmelas

 (Gilbert) Eosebastes
 (Jordan) Eosebastes
deani Starks
introniger (Gilbert) Eosebastes
 (Eigenmann and Eigenmann) Eosebastes
rupestris (Gilbert) Sebastomus

 (Jordan and Thompson)

 Ayres Hispaniscus
 (Eigenmann and Eigenmann) Hispaniscus
 (Jordan and Gilbert) Hispaniscus
atrorubens (Gilbert) Zalopyr
 (Jordan and Gilbert) Zalopyr
 (Jordan and Hubbs)
 (Matsubara)
 Cuvier and Valenciennes
 Gunther
taczanowskii (Steindachner)
 Barsukov
guentheri (Jordan and Starks)
paradoxus Matsubara
tokionis (Jordan and Starks)

 Matsubara
 Temminck and Schlegel
 (Schmidt)
 Matsubara

 Hilgendorf
 Steindachner and Doderlein

ijimae (Jordan and Metz)
zonatus Chen and Barsukov

 (Jordan and Gilbert) Pteropodus
 Richardson Pteropodus
 (Jordan and Gilbert) Pteropodus
gilberti Cramer Pteropodus
 (Jordan and Gilbert) Pteropodus
 Ayres Pteropodus
 (Jordan and Gilbert) Pteropodus
vexillaris (Jordan and Gilbert) Pteropodus
 Hilgendorf
 Hilgendorf
 (Matsubara)
 (Matsubara)

alexandri (Evermann and Goldsborough)
 (Thompson)
 (Jordan and Gilbert) Rosicola
 (Gill) Rosicola

 (Travin)
 Storer
norvegica (Ascanius)
 Kroyer

 Ayres Sebastichthys

albo-fasciatus (Lacepede)
 (Cuvier and Valenciennes)
tertius (Barsukov and Chen)

 (Jordan and Gilbert) Sebastichthys

 (Eigenmann and Eigenmann) Sebastodes
 (Gilbert) Sebastodes
 Ayres Sebastodes
 (Jordan and Starks)
 Hilgendorf

 Chen
 Chen
 Chen
 Chen
 Chen
 Chen
 Ayres
 Chen
ayresii (Gilbert and Cramer) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Eigenmann and Eigenmann) Sebastomus
gillii (Eigenmann) Sebastomus
rhodochloris (Jordan and Gilbert) Sebastomus
 Girard Sebastomus
 (Jordan and Gilbert) Sebastomus
 Valenciennes
 (Gmelin)
chamaco (Evermann and Radcliffe)

 (Tilesius) Primospina
 (Ayres) Sebastosomus
 Girard Sebastosomus
 (Jordan and Gilbert) Primospina
 (Eigenmann and Eigenmann) Sebastosomus

 (Jordan and Gilbert) Sebastomus

 Gunther

 Barsukov
 (Jordan and Evermann) Zalopyr
kawaradai (Matsubara)
melanostictus (Matsubara)
swifti (Evermann and Goldsborough)
 Hilgendorf

 Majorgeneric or
 subgeneric
 assignments

 Jordan,
 Evermann,
 and Clark,
Original species (1) Authors 1930

 (Fitch)
 Lea and Fitch
 (Westrheim and Tsuyuki)
 Barsukov
 (Taranetz and Moiseev)
 Kim and Lee
 Eitner et al.

alutum (Gilbert) Acutomentum
brevispine (Bean) Acutomentum
clavilatum Starks Acutomentum
eigenmanni Cramer Acutomentum
entomelas (Jordan and Gilbert) Acutomentum
 (Cramer) Acutomentum
 (Eigenmann and Beeson) Acutomentum
ovale (Ayres) Acutomentum
rufum (Eigenmann and Eigenmann) Acutomentum
 (Jordan and Starks)
 (Jordan and Starks)
 (Jordan and Snyder)
 (Wakiya)
paucispinosus Matsubara

 Chen
 Chen
 (Beebe and Tee-Van)
 (Gilbert) Hispaniscus
 Quast
 Lea and Fitch
 (Gilbert) Eosebastes
 (Starks) Eosebastes
prorigerum (Jordan and Gilbert) Acutomentum
 (Gilbert) Eosebastes
 (Gilbert) Eosebastes
 (Gilbert) Acutornentum
 (Gilbert) Hispaniscus

dallii (Eigenmann and Beeson) Auctospina
auriculata Girard Auctospina

 Hilgendorf Emmelas

 (Gilbert) Eosebastes
 (Jordan) Eosebastes
deani Starks Eosebastes
introniger (Gilbert) Eosebastes
 (Eigenmann and Eigenmann) Eosebastes
rupestris (Gilbert) Sebastomus

 (Jordan and Thompson)

 Ayres Hispaniscus
 (Eigenmann and Eigenmann) Hispaniscus
 (Jordan and Gilbert) Hispaniscus
atrorubens (Gilbert) Zalopyr
 (Jordan and Gilbert) Zalopyr
 (Jordan and Hubbs)
 (Matsubara)
 Cuvier and Valenciennes
 Gunther
taczanowskii (Steindachner)
 Barsukov
guentheri (Jordan and Starks)
paradoxus Matsubara
tokionis (Jordan and Starks)

 Matsubara
 Temminck and Schlegel
 (Schmidt)
 Matsubara

 Hilgendorf
 Steindachner and Doderlein

ijimae (Jordan and Metz)
zonatus Chen and Barsukov

 (Jordan and Gilbert) Pteropodus
 Richardson Pteropodus
 (Jordan and Gilbert) Pteropodus
gilberti Cramer Pteropodus
 (Jordan and Gilbert) Pteropodus
 Ayres Pteropodus
 (Jordan and Gilbert) Pteropodus
vexillaris (Jordan and Gilbert) Pteropodus
 Hilgendorf
 Hilgendorf
 (Matsubara)
 (Matsubara)

alexandri (Evermann and Goldsborough) Rosicola
 (Thompson) Rosicola
 (Jordan and Gilbert) Rosicola
 (Gill) Rosicola

 (Travin)
 Storer
norvegica (Ascanius) Sebastes
 Kroyer Sebastes

 Ayres Sebastichthys

albo-fasciatus (Lacepede)
 (Cuvier and Valenciennes)
tertius (Barsukov and Chen)

 (Jordan and Gilbert) Sebastocarus

 (Eigenmann and Eigenmann) Sebastodes
 (Gilbert) Sebastodes
 Ayres Sebastodes
 (Jordan and Starks)
 Hilgendorf

 Chen
 Chen
 Chen
 Chen
 Chen
 Chen
 Ayres
 Chen
ayresii (Gilbert and Cramer) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Eigenmann and Eigenmann) Sebastomus
gillii (Eigenmann) Sebastomus
rhodochloris (Jordan and Gilbert) Sebastomus
 Girard Sebastomus
 (Jordan and Gilbert) Sebastomus
 Valenciennes
 (Gmelin)
chamaco (Evermann and Radcliffe)

 (Tilesius) Sebastosomus
 (Ayres) Sebastosomus
 Girard Sebastosomus
 (Jordan and Gilbert) Sebastosomus
 (Eigenmann and Eigenmann) Sebastosomus

 (Jordan and Gilbert) Sebastopyr

 Gunther

 Barsukov
 (Jordan and Evermann) Zalopyr
kawaradai (Matsubara)
melanostictus (Matsubara) Eosebastes
swifti (Evermann and Goldsborough)
 Hilgendorf

 Majorgeneric or
 subgeneric
 assignments

 Matsubara,
Original species (1) Authors 1943

 (Fitch)
 Lea and Fitch
 (Westrheim and Tsuyuki)
 Barsukov
 (Taranetz and Moiseev)
 Kim and Lee
 Eitner et al.

alutum (Gilbert) Acutomentum
brevispine (Bean)
clavilatum Starks
eigenmanni Cramer
entomelas (Jordan and Gilbert)
 (Cramer)
 (Eigenmann and Beeson)
ovale (Ayres)
rufum (Eigenmann and Eigenmann)
 (Jordan and Starks) Acutomentum
 (Jordan and Starks) Acutomentum
 (Jordan and Snyder) Acutomentum
 (Wakiya) Acutomentum
paucispinosus Matsubara Acutomentum

 Chen
 Chen
 (Beebe and Tee-Van)
 (Gilbert)
 Quast
 Lea and Fitch
 (Gilbert)
 (Starks)
prorigerum (Jordan and Gilbert)
 (Gilbert)
 (Gilbert)
 (Gilbert)
 (Gilbert)

dallii (Eigenmann and Beeson)
auriculata Girard

 Hilgendorf Emmelas

 (Gilbert)
 (Jordan)
deani Starks
introniger (Gilbert)
 (Eigenmann and Eigenmann)
rupestris (Gilbert)

 (Jordan and Thompson) Hatumeus

 Ayres
 (Eigenmann and Eigenmann)
 (Jordan and Gilbert)
atrorubens (Gilbert)
 (Jordan and Gilbert)
 (Jordan and Hubbs) Mebarus
 (Matsubara) Mebarus
 Cuvier and Valenciennes Mebarus
 Gunther Mebarus
taczanowskii (Steindachner) Mebarus
 Barsukov
guentheri (Jordan and Starks) Mebarus
paradoxus Matsubara Mebarus
tokionis (Jordan and Starks) Mebarus

 Matsubara Murasoius
 Temminck and Schlegel Murasoius
 (Schmidt) Murasoius
 Matsubara Murasoius

 Hilgendorf Neohispaniscus
 Steindachner and Doderlein Neohispaniscus

ijimae (Jordan and Metz) Neohispaniscus
zonatus Chen and Barsukov

 (Jordan and Gilbert)
 Richardson
 (Jordan and Gilbert)
gilberti Cramer
 (Jordan and Gilbert)
 Ayres
 (Jordan and Gilbert)
vexillaris (Jordan and Gilbert)
 Hilgendorf Pteropodus
 Hilgendorf Pteropodus
 (Matsubara) Pteropodus
 (Matsubara) Pteropodus

alexandri (Evermann and Goldsborough)
 (Thompson)
 (Jordan and Gilbert)
 (Gill)

 (Travin)
 Storer
norvegica (Ascanius)
 Kroyer

 Ayres

albo-fasciatus (Lacepede)
 (Cuvier and Valenciennes)
tertius (Barsukov and Chen)

 (Jordan and Gilbert)

 (Eigenmann and Eigenmann)
 (Gilbert)
 Ayres
 (Jordan and Starks) Sebastodes
 Hilgendorf Sebastodes

 Chen
 Chen
 Chen
 Chen
 Chen
 Chen
 Ayres
 Chen
ayresii (Gilbert and Cramer)
 (Jordan and Gilbert)
 (Jordan and Gilbert)
 (Eigenmann and Eigenmann)
gillii (Eigenmann)
rhodochloris (Jordan and Gilbert)
 Girard
 (Jordan and Gilbert)
 Valenciennes
 (Gmelin)
chamaco (Evermann and Radcliffe)

 (Tilesius)
 (Ayres)
 Girard
 (Jordan and Gilbert)
 (Eigenmann and Eigenmann)

 (Jordan and Gilbert)

 Gunther Takenokius

 Barsukov
 (Jordan and Evermann)
kawaradai (Matsubara) Zalopyr
melanostictus (Matsubara) Zalopyr
swifti (Evermann and Goldsborough)
 Hilgendorf Zalopyr

 Other
 subgeneric
 assignments

Original species (1) Authors Reference

 (Fitch)
 Lea and Fitch
 (Westrheim and Tsuyuki)
 Barsukov
 (Taranetz and Moiseev)
 Kim and Lee
 Eitner et al.

alutum (Gilbert)
brevispine (Bean)
clavilatum Starks
eigenmanni Cramer
entomelas (Jordan and Gilbert) Chen, 1986
 (Cramer) Chen, 1986
 (Eigenmann and Beeson)
ovale (Ayres) Chen, 1986
rufum (Eigenmann and Eigenmann) Chen, 1986
 (Jordan and Starks)
 (Jordan and Starks)
 (Jordan and Snyder)
 (Wakiya)
paucispinosus Matsubara

 Chen Chen, 1986
 Chen Chen, 1986
 (Beebe and Tee-Van) Chen, 1986
 (Gilbert) Hubbs, 1951;
 Chen, 1986
 Quast Chen, 1986
 Lea and Fitch
 (Gilbert) Chen, 1986
 (Starks) Chen, 1986
prorigerum (Jordan and Gilbert) Chen, 1986
 (Gilbert) Chen, 1986
 (Gilbert) Chen, 1986
 (Gilbert) Chen, 1986
 (Gilbert) Chen, 1986

dallii (Eigenmann and Beeson)
auriculata Girard

 Hilgendorf

 (Gilbert)
 (Jordan)
deani Starks
introniger (Gilbert)
 (Eigenmann and Eigenmann)
rupestris (Gilbert)

 (Jordan and Thompson)

 Ayres
 (Eigenmann and Eigenmann)
 (Jordan and Gilbert)
atrorubens (Gilbert)
 (Jordan and Gilbert) Chen, 1985; 1986
 (Jordan and Hubbs) Chen, 1985; 1986
 (Matsubara) Chen, 1985
 Cuvier and Valenciennes Chen, 1985; 1986
 Gunther Chen, 1985; 1986
taczanowskii (Steindachner) Chen, 1985
 Barsukov
guentheri (Jordan and Starks)
paradoxus Matsubara
tokionis (Jordan and Starks)

 Matsubara
 Temminck and Schlegel
 (Schmidt)
 Matsubara

 Hilgendorf
 Steindachner and Doderlein Chen, 1986

ijimae (Jordan and Metz) Chen, 1986
zonatus Chen and Barsukov Chen, 1986

 (Jordan and Gilbert)
 Richardson
 (Jordan and Gilbert)
gilberti Cramer
 (Jordan and Gilbert)
 Ayres
 (Jordan and Gilbert)
vexillaris (Jordan and Gilbert)
 Hilgendorf Chen, 1986
 Hilgendorf Chen, 1986
 (Matsubara) Jordan and
 Hubbs, 1925
 (Matsubara)

alexandri (Evermann and Goldsborough)
 (Thompson)
 (Jordan and Gilbert)
 (Gill)

 (Travin)
 Storer
norvegica (Ascanius)
 Kroyer

 Ayres

albo-fasciatus (Lacepede) Matsubara, 1943
 (Cuvier and Valenciennes) Matsubara, 1943
tertius (Barsukov and Chen) Chen, 1986

 (Jordan and Gilbert)

 (Eigenmann and Eigenmann)
 (Gilbert)
 Ayres
 (Jordan and Starks)
 Hilgendorf

 Chen Chen, 1975
 Chen Chen, 1971
 Chen Chen, 1971
 Chen Chen, 1971
 Chen Chen, 1971
 Chen Chen, 1971
 Ayres
 Chen Chen, 1971
ayresii (Gilbert and Cramer)
 (Jordan and Gilbert)
 (Jordan and Gilbert)
 (Eigenmann and Eigenmann)
gillii (Eigenmann)
rhodochloris (Jordan and Gilbert)
 Girard
 (Jordan and Gilbert)
 Valenciennes Chen, 1971
 (Gmelin) Chen, 1971
chamaco (Evermann and Radcliffe) Chen, 1971

 (Tilesius)
 (Ayres)
 Girard
 (Jordan and Gilbert)
 (Eigenmann and Eigenmann)

 (Jordan and Gilbert)

 Gunther Chen, 1986

 Barsukov
 (Jordan and Evermann)
kawaradai (Matsubara)
melanostictus (Matsubara)
swifti (Evermann and Goldsborough)
 Hilgendorf

 Other
 subgeneric
 assignments

Original species (1) Authors Subgenus

 (Fitch) Unknown
 Lea and Fitch Unknown
 (Westrheim and Tsuyuki) Unknown
 Barsukov Unknown
 (Taranetz and Moiseev) Unknown
 Kim and Lee Unknown
 Eitner et al. Unknown

alutum (Gilbert)
brevispine (Bean)
clavilatum Starks
eigenmanni Cramer
entomelas (Jordan and Gilbert) Acutomentum
 (Cramer) Acutomentum
 (Eigenmann and Beeson)
ovale (Ayres) Acutomentum
rufum (Eigenmann and Eigenmann) Acutomentum
 (Jordan and Starks)
 (Jordan and Starks)
 (Jordan and Snyder)
 (Wakiya)
paucispinosus Matsubara

 Chen Allosebastes
 Chen Allosebastes
 (Beebe and Tee-Van) Allosebastes
 (Gilbert) Allosebastes
 Quast Allosebastes
 Lea and Fitch
 (Gilbert) Allosebastes
 (Starks) Allosebastes
prorigerum (Jordan and Gilbert) Allosebastes
 (Gilbert) Allosebastes
 (Gilbert) Allosebastes
 (Gilbert) Allosebastes
 (Gilbert) Allosebastes

dallii (Eigenmann and Beeson)
auriculata Girard

 Hilgendorf

 (Gilbert)
 (Jordan)
deani Starks
introniger (Gilbert)
 (Eigenmann and Eigenmann)
rupestris (Gilbert)

 (Jordan and Thompson)

 Ayres
 (Eigenmann and Eigenmann)
 (Jordan and Gilbert)
atrorubens (Gilbert)
 (Jordan and Gilbert) Mebarus
 (Jordan and Hubbs) Mebarus
 (Matsubara) not Mebarus
 Cuvier and Valenciennes Mebarus
 Gunther Mebarus
taczanowskii (Steindachner) not Mebarus
 Barsukov
guentheri (Jordan and Starks)
paradoxus Matsubara
tokionis (Jordan and Starks)

 Matsubara
 Temminck and Schlegel
 (Schmidt)
 Matsubara

 Hilgendorf
 Steindachner and Doderlein Takenokius

ijimae (Jordan and Metz) Takenokius
zonatus Chen and Barsukov Takenokius

 (Jordan and Gilbert)
 Richardson
 (Jordan and Gilbert)
gilberti Cramer
 (Jordan and Gilbert)
 Ayres
 (Jordan and Gilbert)
vexillaris (Jordan and Gilbert)
 Hilgendorf Takenokius
 Hilgendorf Takenokius
 (Matsubara) Sebastocles
 (Matsubara)

alexandri (Evermann and Goldsborough)
 (Thompson)
 (Jordan and Gilbert)
 (Gill)

 (Travin)
 Storer
norvegica (Ascanius)
 Kroyer

 Ayres

albo-fasciatus (Lacepede) Sebastiscus
 (Cuvier and Valenciennes) Sebastiscus
tertius (Barsukov and Chen) Sebastiscus

 (Jordan and Gilbert)

 (Eigenmann and Eigenmann)
 (Gilbert)
 Ayres
 (Jordan and Starks)
 Hilgendorf

 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Ayres
 Chen Sebastomus
ayresii (Gilbert and Cramer)
 (Jordan and Gilbert)
 (Jordan and Gilbert)
 (Eigenmann and Eigenmann)
gillii (Eigenmann)
rhodochloris (Jordan and Gilbert)
 Girard
 (Jordan and Gilbert)
 Valenciennes Sebastomus
 (Gmelin) Sebastomus
chamaco (Evermann and Radcliffe) Sebastomus

 (Tilesius)
 (Ayres)
 Girard
 (Jordan and Gilbert)
 (Eigenmann and Eigenmann)

 (Jordan and Gilbert)

 Gunther Takenokius

 Barsukov
 (Jordan and Evermann)
kawaradai (Matsubara)
melanostictus (Matsubara)
swifti (Evermann and Goldsborough)
 Hilgendorf
 Current
Original species (1) Authors subgenus

 (Fitch) Unknown
 Lea and Fitch Unknown
 (Westrheim and Tsuyuki) Unknown
 Barsukov Unknown
 (Taranetz and Moiseev) Unknown
 Kim and Lee Unknown
 Eitner et al. Unknown

alutum (Gilbert) Acutomentum
brevispine (Bean) Acutomentum
clavilatum Starks Acutomentum
eigenmanni Cramer Acutomentum
entomelas (Jordan and Gilbert) Acutomentum
 (Cramer) Acutomentum
 (Eigenmann and Beeson) Acutomentum
ovale (Ayres) Acutomentum
rufum (Eigenmann and Eigenmann) Acutomentum
 (Jordan and Starks) Acutomentum
 (Jordan and Starks) Acutomentum
 (Jordan and Snyder) Acutomentum
 (Wakiya) Acutomentum
paucispinosus Matsubara Acutomentum

 Chen Allosebastes
 Chen Allosebastes
 (Beebe and Tee-Van) Allosebastes
 (Gilbert) Allosebastes
 Quast Allosebastes
 Lea and Fitch Allosebastes
 (Gilbert) Allosebastes
 (Starks) Allosebastes
prorigerum (Jordan and Gilbert) Allosebastes
 (Gilbert) Allosebastes
 (Gilbert) Allosebastes
 (Gilbert) Allosebastes
 (Gilbert) Allosebastes

dallii (Eigenmann and Beeson) Auctospina
auriculata Girard Auctospina

 Hilgendorf Emmelas

 (Gilbert) Eosebastes
 (Jordan) Eosebastes
deani Starks Eosebastes
introniger (Gilbert) Eosebastes
 (Eigenmann and Eigenmann) Eosebastes
rupestris (Gilbert) Eosebastes

 (Jordan and Thompson) Hatumeus

 Ayres Hispaniscus
 (Eigenmann and Eigenmann) Hispaniscus
 (Jordan and Gilbert) Hispaniscus
atrorubens (Gilbert) Mebarus
 (Jordan and Gilbert) Mebarus
 (Jordan and Hubbs) Mebarus
 (Matsubara) Mebarus
 Cuvier and Valenciennes Mebarus
 Gunther Mebarus
taczanowskii (Steindachner) Mebarus
 Barsukov Mebarus
guentheri (Jordan and Starks) Mebarus
paradoxus Matsubara Mebarus
tokionis (Jordan and Starks) Mebarus

 Matsubara Murasoius
 Temminck and Schlegel Murasoius
 (Schmidt) Murasoius
 Matsubara Murasoius

 Hilgendorf Neohispaniscus
 Steindachner and Doderlein Neohispaniscus

ijimae (Jordan and Metz) Neohispaniscus
zonatus Chen and Barsukov Neohispaniscus

 (Jordan and Gilbert) Pteropodus
 Richardson Pteropodus
 (Jordan and Gilbert) Pteropodus
gilberti Cramer Pteropodus
 (Jordan and Gilbert) Pteropodus
 Ayres Pteropodus
 (Jordan and Gilbert) Pteropodus
vexillaris (Jordan and Gilbert) Pteropodus
 Hilgendorf Pteropodus
 Hilgendorf Pteropodus
 (Matsubara) Pteropodus
 (Matsubara) Pteropodus

alexandri (Evermann and Goldsborough) Rosicola
 (Thompson) Rosicola
 (Jordan and Gilbert) Rosicola
 (Gill) Rosicola

 (Travin) Sebastes
 Storer Sebastes
norvegica (Ascanius) Sebastes
 Kroyer Sebastes

 Ayres Sebastichthys

albo-fasciatus (Lacepede) Sebastiscus
 (Cuvier and Valenciennes) Sebastiscus
tertius (Barsukov and Chen) Sebastiscus

 (Jordan and Gilbert) Sebastocarus

 (Eigenmann and Eigenmann) Sebastodes
 (Gilbert) Sebastodes
 Ayres Sebastodes
 (Jordan and Starks) Sebastodes
 Hilgendorf Sebastodes

 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Chen Sebastomus
 Ayres Sebastomus
 Chen Sebastomus
ayresii (Gilbert and Cramer) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Jordan and Gilbert) Sebastomus
 (Eigenmann and Eigenmann) Sebastomus
gillii (Eigenmann) Sebastomus
rhodochloris (Jordan and Gilbert) Sebastomus
 Girard Sebastomus
 (Jordan and Gilbert) Sebastomus
 Valenciennes Sebastomus
 (Gmelin) Sebastomus
chamaco (Evermann and Radcliffe) Sebastomus

 (Tilesius) Sebastosomus
 (Ayres) Sebastosomus
 Girard Sebastosomus
 (Jordan and Gilbert) Sebastosomus
 (Eigenmann and Eigenmann) Sebastosomus

 (Jordan and Gilbert) Sebastropyr

 Gunther Takenokius

 Barsukov Zalopyr
 (Jordan and Evermann) Zalopyr
kawaradai (Matsubara) Zalopyr
melanostictus (Matsubara) Zalopyr
swifti (Evermann and Goldsborough) Zalopyr
 Hilgendorf Zalopyr

(1) Listed when different from present species name.

(2) () = synonomyzed with; * = form of pachycephalus.
Table 1.--Sebastiscus, and Hozukius, and Sebastes subgenera: type
species, ranges, numbers of species, and authors.

 No. of
Subgenus Type species Range species

Acutomentum ovalis NE-NW Pac. 11
Allosebastes sinensis NE Pac.-Gulf Calif. 14
Auctospina auriculatus NE Pac. 2
Emmelas glaucus NW Pac. 1
Eosebastes aurora NE Pac. 3
Hatumeus owstoni NW Pac. 1
Hispaniscus rubrivinctus NE Pac. 3
Hozukius emblemarius (1) NW Pac. 2
Mebarus inermis NE-NW Pac. 7
Murasoius pachycephalus NW Pac. 1
Neohispaniscus schlegeli NW Pac. 2
Primospina mystinus NE Pac. 1
Pteropodus maliger NE-NW Pac. 10
Rosicola pinniger NE Pac. 3
Sebastes norvegicus N Atl. 4
Sebastichthys nigrocinctus NE Pac. 1
Sebastiscus marmoratus NW Pac. 2
Sebastocarus serriceps NE Pac. 1
Sebastocles hubbsi NW Pac. 1
Sebastodes paucispinis NE-NW Pac. 5
Sebastomus rosaceus NE Pac.-S Hemis. 16
Sebastopyr rubberimus NE Pac. 1
Sebastosomus melanops NE Pac. 5
Takenokius oblongus NW Pac. 1
Zalopyr aleutianus NE-NW Pac. 3

Subgenus Authors

Acutomentum Eigenmann and Beeson, 1893
Allosebastes Hubbs, 1951
Auctospina Eigenmann and Beeson, 1893
Emmelas Jordan and Evermann, 1898
Eosebastes Jordan and Evermann, 1896
Hatumeus Matsubara, 1943
Hispaniscus Jordan and Evermann, 1896
Hozukius Matsubara, 1934
Mebarus Matsubara, 1943
Murasoius Matsubara, 1943
Neohispaniscus Matsubara, 1943
Primospina Eigenmann and Beeson, 1893
Pteropodus Eigenmann and Beeson, 1893
Rosicola Jordan and Evermann, 1896
Sebastes Ascanius, 1772
Sebastichthys Gill, 1862
Sebastiscus Jordan and Starks, 1904
Sebastocarus Jordan and Evermann, 1927
Sebastocles Jordan and Hubbs, 1925
Sebastodes Gill, 1861
Sebastomus Gill, 1864
Sebastopyr Jordan and Evermann, 1927
Sebastosomus Gill, 1864
Takenokius Matsubara, 1943
Zalopyr Jordan and Evermann, 1898

Subgenus Reference

Acutomentum Am. Nat. 27:669
Allosebastes Proc. Biol. Soc. Wash. 64:129-130
Auctospina Am. Nat. 27:670
Emmelas Bull. U.S. Natl. Mus. 47:1773
Eosebastes Rep. U.S. Comm. Fish Fish. 1895, pt. 21:430
Hatumeus Trans. Sigenkagaku Kenkyusyo
Hispaniscus Rep. U.S. Comm. Fish Fish, 1895, pt. 21:431
Hozukius J. Imp. Fish. Inst. Tokyo 30:199-210.
Mebarus Trans. Sigenkagaku Kenkyusyo:194
Murasoius Trans. Sigenkagaku Kenkyusyo:235
Neohispaniscus Trans. Sigenkagaku Kenkyusyo:226
Primospina Am. Nat. 27:669
Pteropodus Am. Nat. 27:670
Rosicola Rep. U.S. Comm. Fish Fish. 1895, pt. 21:429
Sebastes Claude Philibert, pt. 2
Sebastichthys Proc. Acad. Nat. Sci. Phila. 14:278, 329
Sebastiscus Proc. U.S. Nat. Mus. 27:91-175
Sebastocarus Proc. Calif. Acad. Sci., Ser. 4, 16:507
Sebastocles Mem. Carnegie Mus. 10:260.
Sebastodes Proc. Acad. Nat. Sci. Phila. 13:165
Sebastomus Proc. Acad. Nat. Sci. Phila. 16:147
Sebastopyr Proc. Calif. Acad. Sci., Ser. 4, 16:506
Sebastosomus Proc. Acad. Nat. Sci. Phila. 16:147
Takenokius Trans. Sigenkagaku Kenkyusyo:233
Zalopyr Bull. U.S. Natl. Mus. 47:1795

(1) The species name emblemarius has been misspelled embremarius in
several publications including Matsubara (1934, 1943).
Table 2.--Major field guides to identify species of Northeast Pacific
Sebastes.

Reference Geographic area

Clemens and Wilby (1946) British Columbia.

Phillips (1957) California, but ranges of some species
 extend to Baja California and Alaska.

Clemens and Wilby (1961) British Columbia.

Hitz (1965) (1) Baja California to the Bering Sea.

Miller and Lea (1972) California, but ranges of some species
 extend to Baja California and Alaska.

Hart (1973) British Columbia.

Eschmeyer et al. (1983) Coastal waters from the Gulf of Alaska
 to Baja California.

Kramer and O'Connell (1988) Species that occur off Alaska, includes
 most species from British Columbia,
 Washington, and Oregon also.

Orr et al. (2000) Northeast Pacific Ocean north of Mexico,
 including the eastern Bering Sea.

Reference No. of species

Clemens and Wilby (1946) 20

Phillips (1957) 49

Clemens and Wilby (1961) 23

Hitz (1965) (1) 53

Miller and Lea (1972) 58

Hart (1973) 33

Eschmeyer et al. (1983) 58

Kramer and O'Connell (1988) 35

Orr et al. (2000) 66

Reference Presentation

Clemens and Wilby (1946) Extended individual species accounts
 including airbrush monochrome drawings.

Phillips (1957) Key to the species and individual species
 accounts with monochrome photographs.

Clemens and Wilby (1961) Revision of Clemens and Wilby (1946).

Hitz (1965) (1) 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) Brief individual species accounts and a
 key illustrated with monochrome sketches
 showing important characters.

Hart (1973) Key to the species and extended
 individual species accounts with
 monochrome drawings.

Eschmeyer et al. (1983) Individual species accounts and color
 plates annotated with important
 characters.

Kramer and O'Connell (1988) Brief individual species accounts
 including head spine diagrams, sketches
 of important characters, and color
 photographs.

Orr et al. (2000) Format follows Hitz (1965). Includes
 individual species accounts and
 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)

 3 A1 fibers long
17 insertion of transversus branched
 dorsalis anterior
41 swimbladder present
56 infraorbitals 4 & 5 both present
64 pharyngobranchials 2 & 3, all separate
 upper pharyngeal
70 number of vertebrae 24
80 number of hypurals 2+3
83 scapula and upper radial autogenous
88 supraneural 1 present

 Characters States and coding

Number Description Intermediate (1) Derived (2)

 3 A1 fibers short
17 insertion of transversus unbranched
 dorsalis anterior
41 swimbladder absent
56 infraorbitals 4 & 5 one present both absent
64 pharyngobranchials 2 & 3, pharyngobranchials all fused
 upper pharyngeal fused
70 number of vertebrae more than 24
80 number of hypurals 1+3 1+2
83 scapula and upper radial fused
88 supraneural absent

 Characters

Number Description Distribution of derived states
 outside Sebastidae

 3 A1 fibers Congiopodidae
17 insertion of transversus Congiopodidae and occasional
 dorsalis anterior elsewhere
41 swimbladder Widespread, especially
 Scorpaenidae
56 infraorbitals 4 & 5 Very widespread
64 pharyngobranchials 2 & 3, Intermediate rare, derived
 upper pharyngeal Aploactinidae and Pataecidae
70 number of vertebrae Widespread except Setarchidae and
 Scorpaenidae
80 number of hypurals Derived states widespread
83 scapula and upper radial Widespread
88 supraneural Very widespread
Table 4.--Characters and character states for analysis
of Sebastidae, following Ishida, 1994.

 Characters (1)

Genus 3 17 41 56 64 70 80 83 88

Sebastes 1 0 0 0 0 1 2 1 0
Sebastiscus 1 0 0 0 0 1 2 1 0
Hozukius 1 0 0 0 0 1 2 1 0
Helicolenus 1 0 1 0 0 1 2 1 0
Adelosebastes 1 0 1 0 0 1 2 1 0
Sebastolobus 1 0 1 0 0 1 2 0 0
Trachyscorpia 1 0 1 1 0 1 0 0 0
Plectrogenium 1 1 1 1 1 1 0 0 1

(1) 0=primitive, 1=intermediate, 2=derived.
Table 5.--Ecological correlated of characters of Sebastes.

 Demersal Pelagic
Character state state

Head spines strength Strong Weak
Head spine numbers Many Few
Jaw length Lower > upper Equal
Interorbital shape Concave Convex
Lower pectoral rays Thick Not thick
Gill raker length Short Long
Jaws scaled Yes No
Base of skull Curved Flat
Mesethmoid process Strong Weak
Color Red Black

 Low latitude High latitude
 state state

Vertebrae <29 >28
Dorsal spines <14 14-15
Dorsal rays <14 15-16
Anal rays 5 9
Principal caudal rays 13 15
Pectoral rays 17-18 >18


Acknowledgments

I am pleased to express appreciation to the following people for their help with this work. Bruce Collette, NMFS Systematics Laboratory, supplied draft copies of work in progress on North Atlantic rockfishes and rockfish specimens from the North Atlantic for study. Abby J. Simpson, formerly of the School of Fisheries, University of Washington, Seattle, kindly translated Cuvier's original description of Sebastes from French. Tony Gharrett, University of Alaska; Geoff Moser, SWFSC; and Jay Orr, AFSC, lent much needed encouragement for this project and reviewed an earlier draft of the manuscript. Mike Canino, AFSC, and Paul Bentzen, University of Washington, reviewed an earlier draft also.

(1) Sebastiscus and Hozukius have been considered subgenera of Sebastes by some and separate but closely related genera by others. This paper will consider each a separate genus and will focus primarily on the taxonomic history of Sebastes. Sebastes is by far the most species-rich and widely distributed genus; Sebastiscus and Hozukius have three and two species, respectively, and occur only in the northwest Pacific. As will be discussed later, Sebastes and these other two genera form an unresolved trichotomy within the Sebastidae of the suborder Scorpaenoidei (Ishida, 1994). These three genera could be given the subfamily name Sebastinae, following Matsubara (1943).

(2) See Simpson, A. J., A. W. Kendall, Jr., and J. W. Orr. 2000. Translations of the original descriptions of Sebastes by Georges Cuvier. U.S. Dep. Commer., NOAA, Natl. Mar. Fish. Serv., Alaska Fish. Sci. Cent., 7600 Sand Point Way NE, Seattle WA 98115. Proc. Rep. 2000-09, 24 p.

(3) Sebastes auriculatus is in the subgenus Auctospina with S. dalli. However, this study and others (e.g. Johnson et al., 1972) indicated it is closely related to other shallow-water deep-bodied spiny rockfishes (e.g. S. caurinus) that are in Pteropodus.

(4) Ida, Hitoshi, Kitasato Univ., Sanriku, Jpn., personal commun., 20 January 1999.

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Arthur W. Kendall, Jr., retired 3 January 2001, was with the Alaska Fisheries Science Center, National Marine Fisheries Service, NOAA, 7600 Sand Point Way N.E., Seattle, WA 98115.
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Title Annotation:rockfish research
Author:Kendall, Arthur W., Jr.
Publication:Marine Fisheries Review
Date:Mar 22, 2000
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