Revision of the genus Lepidopsetta Gill, 1862 based on larval and adult morphology, with a description of a new species from the North Pacific Ocean and Bering Sea – Teleostei: Pleuronectidae

Revision of the genus Lepidopsetta Gill, 1862 based on larval and adult morphology, with a description of a new species from the North Pacific Ocean and Bering Sea – Teleostei: Pleuronectidae – Statistical Data Included

James W. Orr

Abstract.–The endemic North Pacific pleuronectid genus Lepidopsetta Gill is revised to include three species: L. bilineata (Ayres), L. polyxystra n. sp., and L. mochigarei Snyder. Adults of L. bilineata can be distinguished by a low gill-raker count and high supraorbital pore count; larvae may be distinguished by four dorsal midline melanophores, heavy finfold pigment, a short snout-to-anus length, and a deep body. The species ranges from Baja California to the eastern Aleutian Islands and the extreme southeastern Bering Sea. Adults of L. mochigarei are distinguished from all other members of Lepidopsetta by higher scale and preopercular pore counts and lower gill-raker and supraorbital pore counts. Larvae are similar to larvae of L. bilineata but can be distinguished by their postanal pigment pattern and melanophores on pectoral-fin rays. Lepidopsetta mochigarei ranges from the southern Sea of Okhotsk to Korea. Adults of L. polyxystra n. sp. are diagnosed by a high gill-raker count and low supraorbital pore count; larvae are diagnosed by two dorsal midline melanophores, light finfold pigment, long snout-to-anus length, and a slender body. The species is found from Puget Sound through the Bering Sea and Aleutian Islands to the Kuril Islands, overlapping with L. bilineata from the extreme southeastern Bering Sea to Puget Sound and with L. mochigarei in the southern Sea of Okhotsk. Synonymies, diagnoses, descriptions, and geographic distributions are provided for adults and larvae of all species; keys are provided for adults. Descriptions of early juveniles of eastern North Pacific species are also presented. Under the name of L. bilineata, L. polyxystra n. sp. has been the subject of many studies of eastern North Pacific Lepidopsetta. All previous studies of specimens from the southeastern Bering Sea into Puget Sound should be considered applicable at the generic level only, unless voucher specimens are verifiable.

The flatfishes (Pleuronectiformes) of the eastern North Pacific Ocean constitute a major component of the commercial fisheries of the region. In the Bering Sea, which encompasses the largest fisheries resource of the United States, the rock soles of the genus Lepidopsetta are the second most abundant flatfishes and the third most abundant commercial groundfish species, second only to yellowfin sole (Limanda aspera) and walleye pollock (Theragra chalcogramma) (NMFS, 1999).

At the species level, eastern North Pacific pleuronectids have been considered well known. Nearly all species were recognized and described during the latter half of the 1800s, primarily through the activities of California ichthyologists. Among the names that remain valid, the last species to be described was Limanda sahkalinensis Hubbs, 1915, although in the western Pacific Microstomus shuntovi Borets, 1983, was most recently described from the northwestern Hawaiian ridge. However, these earlier works were based on adult morphology, and only recently has a knowledge of the ontogeny of these species been acquired. Among the descriptions of early life history stages of eastern North Pacific pleuronectids, one morphological form could not be traced to a recognized species. Examination of this form led to the following revision of the genus Lepidopsetta.

Four nominal species have been described and allocated to the genus Lepidopsetta: Platessa bilineata Ayres, 1855a, was described from San Francisco material, and apparently without the knowledge of Ayres’ slightly earlier description, a specimen collected near Puget Sound was described as Platichthys umbrosus Girard, 1856. Gill (1862) erected Lepidopsetta to contain Platichthys umbrosus and later (1864) indicated that Platessa bilineata Ayres, 1855a, was allied and perhaps congeneric. Lockington (1879b) published a redescription of L. umbrosa, describing the misidentified new species Isopsetta isolepis (Lockington, 1880a), which he ultimately removed from Lepidopsetta to his new genus Isopsetta (Lockington, 1883). In his description of Lepidopsetta isolepis, he treated Platichthys umbrosus Girard as a synonym of L. bilineata. Nearly 20 years after Gill’s erection of Lepidopsetta, Cope (1873) described Pleuronectes perarcuatus from Alaska, later considered a synonym of L. bilineata by Jordan and Gilbert (1881). Jordan and Evermann (1898) considered each of these nominal species members of Lepidopsetta and further as synonyms of L. bilineata, although they recognized the northern populations (“Puget Sound and northward”) as L. bilineata umbrosa. Finally, Japanese Lepidopsetta were described by Snyder (1911) as L. mochigarei and Jordan and Hubbs (1925) considered all Japanese Lepidopsetta to be representatives of this species.

More recently, the genus has been envisioned as containing either two species, L. bilineata of the eastern North Pacific Ocean and Bering Sea and L. mochigarei of the western North Pacific around Japan (Hubbs, 1915; Sakamoto, 1984b), or a single species, L. bilineata, with two subspecies L. b. bilineata and L. b. mochigarei in the Bering Sea and western Pacific (Taranets, 1937; Schmidt, 1950). A third subspecies in the southeastern North Pacific was recognized by Moiseev (1953) and Wilimovsky et al. (1967). Although both authors split eastern Pacific populations into two subspecies, Moiseev (1953) considered the northern subspecies to be L. b. bilineata and the southern subspecies to be L. b. umbrosa, whereas Wilimovsky et al. (1967) applied the names L. b. perarcuata and L. b. bilineata to northern and southern populations, respectively.

Our study arose from an examination of collections from ichthyoplankton surveys conducted from the Bering Sea to northern California by the Recruitment Processes Task (referred to simply as “Task” in the following account) of the Alaska Fisheries Science Center (AFSC). In 1985, ichthyoplankton taxonomists began to routinely separate an unidentified pleuronectid from numerous collections. Larvae from the developmental series of this unidentified pleuronectid most closely resembled larvae previously described as Psettichthys melanostictus (Hickman, 1959; Pertseva-Ostroumova, 1961) and, for the following few years, larval pigment patterns and morphological characters were used to separate what were then considered Psettichthys into two readily distinguishable morphotypes. Thus in 1986, an early draft of the Task’s laboratory guide (Matarese et al., 1989) included illustrations of an unidentified series referred to as a variant of Psettichthys (“Psettichthys 2”).

During winter and spring 1987, larvae of both morphotypes of “Psettichthys” were collected from Puget Sound and reared through transformation and settlement stages, and upon examination of the reared juveniles, Kendall and Matarese(1) determined that larvae previously referred to as “Psettichthys 2” were, in fact, another form of Lepidopsetta. From the early results of this work, Matarese et al. (1989) decided to include a partial description of the unknown pleuronectid (referred to as “Lepidopsetta 2”) and compared various stages with larvae of Psettichthys and other Lepidopsetta. Mulligan et al. (1995) finally verified the identity of Lepidopsetta 2 by rearing larvae spawned from Lepidopsetta adults collected in Puget Sound and conducted a morphological study of adults. Although they reported significant heterogeneity in shape, structure, and allozymes, they recommended the retention of Wilimovsky et al.’s (1967) subspecies designations.

Although previous authors (Townsend, 1936, 1937; Wilimovsky et al., 1967; Mulligan et al., 1995) concluded that observed variation represented a cline smoothly grading from California through the Bering Sea to Japan and supported subspecific designations, new data became available in 1992 with the development of a fishery for Lepidopsetta in the northern Gulf of Alaska. Domestic fisheries observers (see “Acknowledgments” section) experienced in sampling in the Bering Sea flatfish fisheries began to report the presence of two syntopic adult forms of Lepidopsetta in the northern Gulf of Alaska, one distinctly different from the form in the eastern Bering Sea. These observations spurred a further examination into the morphological differences of all life stages of Lepidopsetta. Our revision therefore incorporates evidence from adult, juvenile, and larval morphology and distribution to support the recognition of three species, one described as new, in the North Pacific. We describe morphological variation in adults,juveniles, and larvae; differentially diagnose adults, juveniles, and larvae; and describe geographic and bathymetric distributions of the three species.

Materials and methods

Unless indicated otherwise, standard length (SL) is used throughout. Institutional abbreviations follow Leviton et al. (1985) and Leviton and Gibbs (1988), as modified by Poss and Collette (1995), except for the Kamchatka Institute of Ecology, abbreviated as KIE.(2)

Adult morphology

Meristic data, except gill-raker counts, and morphometric data were taken from the ocular-side of adult material, following Hubbs and Lagler (1958) with the following exceptions. Standard, head, and snout length were measured from the anterior margin of the maxilla, with the mouth closed. Body depth was the greatest depth measured at the origin of the anal fin. Head length included only the opercle and not the opercular membrane. Snout length was measured to the anterior edge of the dorsal orbit. Cheek length was the greatest distance from the posterior rim of the ventral orbit to the edge of the preopercle, often the posterior angle of the preopercle. Interorbital width equals the least bony width. Pectoral-fin length was the length of the longest ray, often the third ray, and was measured for both ocular-side and blind-side fins. Lateral-line arch length was the distance between anterior and posterior flexion points, and this straight line was used as the base for the depth measurement. Greatest caudal peduncle depth was measured at the base of the caudal fin.

All rayed elements were included in counts of fin rays. The last two rays of the dorsal and anal fins were counted separately. Scales above the lateral line were counted on a diagonal at the greatest depth between the dorsal-fin base and lateral line. Scales below the lateral line were counted from the anal-fin origin on a diagonal to the lateral line. Total scales above and below the lateral line is the sum of the two counts. Cheek scales were counted at greatest cheek length. Ocular-side suborbital pores were counted from the first pore branching from the temporal canal to the anteriormost pore; all pores were counted, including those dorsal and ventral to the canal. Blind-side suborbital pores were counted from the first pore branching from the postorbital canal to the anteriormost pores. Pore counts of the dorsal anterior and posterior branches of the accessory dorsal branch of the lateral line (ADB) began with the first pore of each branch and did not include the single pore typically found at the intersection of the two branches. Supraorbital pores include the first pore dorsal to the postorbital canal and include those pores at the posterior rim of the dorsal orbit (Fig. 1). Preopercular pores begin with the first pore posterior to the mandibular articulation and end at the last pore before the temporal canal. Caudal-fin terminology follows Sakamoto (1984a).

[Figure 1 ILLUSTRATION OMITTED]

Larval collections

Most data were garnered from samples collected by the AFSC with standard bongo gear during ichthyoplankton surveys of the Bering Sea and Gulf of Alaska (1971-1994, Table 1, Appendix Table 1). In most cases, the identities of specimens collected prior to 1985 (when the unidentified pleuronectid developmental series was first separated) have been verified. All larval specimens used for illustrations and morphological descriptions have been cataloged at the University of Washington Fish Collection (UWFC); other material will be archived at the UWFC as well (see Appendix Table 1). Additional data for Lepidopsetta in Canadian waters were obtained from the Canadian Department of Fisheries and Oceans, Pacific Biological Station, Nanaimo (W. Shaw), and Vancouver Public Aquarium (J. Marliave; these specimens are now deposited at UWFC). The Washington Department of Fisheries also provided distribution and life history data for Puget Sound.(3) In addition, dip-net sampling in Puget Sound conducted by the AFSC from 1985 through 1995 resulted in 16 collections of Lepidopsetta larvae (Busby et al., 2000). Two larvae from the Washington, Oregon, and northern California survey area were examined–the only two Lepidopsetta larvae collected during the eight years (from 1980 to 1987) during which the AFSC conducted ichthyoplankton cruises in the region. Geoff Moser, NMFS Southwest Fisheries Science Center (SWFSC), provided data for Lepidopsetta collected during California Cooperative Fisheries Investigations (CalCOFI) cruises conducted along the coast of California.

Table 1 Collections of Lepidopsetta larvae examined for this study. AFSC = Alaska Fisheries Science Center, Seattle, WA; VPA = Vancouver Public Aquarium, Vancouver, BC; PBS = Pacific Biological Station, Nanaimo, BC; CalCOFI = California Current Oceanic Fisheries Investigations, La Jolla, CA.

Source

General location of data Years

Bering Sea AFSC 1971, 1977, 1979, 1988, 1992-94

Gulf of Alaska AFSC 1972, 1978-79, 1981-94

British Columbia VPA, PBS 1983-88, 1991-93

Puget Sound AFSC 1985-95

WA, OR, and N. CA AFSC 1980-87

California CalCOFI 1951-84

Months No. of

General location sampled samples

Bering Sea Apr-Aug 152

Gulf of Alaska Mar-Oct 1896

British Columbia Mar-July 72

Puget Sound Mar-July 16

WA, OR, and N. CA Apr-May 2

California Feb-July 21

Identification of juveniles

Recently transformed juveniles present particular problems in identification because adult characters, such as meristics and pigment patterns, are not completely developed and some aspects of larval shape, structure, and pigment patterns are retained. Thus, early juveniles (approximately 20-35 mm SL) were identified by a combination of characters, including gill-raker counts of the first arch, larval pigment characters retained on the blind side, and size at transformation. Although gill rakers were not fully formed, specimens with counts greater than seven on the lower arch were assumed to be the new species. Juveniles with counts of seven or less were identified in conjunction with other characters. Postsettlement juveniles greater than 30 mm SL were separated by using the above characters in addition to counts of the number of supraorbital pores. Lepidopsetta bilineata typically has 4-6 head pores along the supraorbital canal, whereas the new species typically has 1-3. Differences in gill-raker structure useful in distinguishing adults are usually not evident at sizes less than 200 mm SL.

Identification of larvae

Identifcation of Lepidopsetta larvae was accomplished by a variety of methods because the more traditional serial approach was not possible. Initially, our unidentified pleuronectid series was grouped together on the basis of a continuous sequence of shared characters. This series, however, could not be linked with a recognized pleuronectid taxon whose larvae had not already been described. The unidentified series (Psettichthys 2) was similar to larvae previously described for P. melanostictus. Larvae of Psettichthys 2 were determined to be Lepidopsetta by rearing wild-caught larvae(1) and finally confirmed to be Lepidopsetta by rearing larvae spawned from Lepidopsetta adults collected in Puget Sound (Mulligan et al., 1995).

Nomenclature of larval developmental stages follows Kendall et al. (1984) where each stage is based on widespread, fundamental features of development. The early life history stages used in our study were based on the flexion of the notochord that accompanies the hypochordal development of the homocercal caudal fin. The onset of the juvenile stage is defined in our study as completion of eye migration and attainment of the adult complement of fin elements. Early juveniles (approximately 20-35 mm SL) are defined as those collected in the water column; many had remnants of larval pigmentation on their blind side. Postsettlement juveniles were collected with bottom gear and rarely retained any larval pigmentation on their blind side.

Only melanistic pigment is described in our study because formalin fails to preserve other pigments (live larvae have yellow, orange, and brown chromatophores associated with the melanophores). Previous descriptions of larval pigment patterns, particularly those of pleuronectids, have used confusing terminology. Postanal pigment patterns have been variously described as vertically oriented bands (Pertseva-Ostroumova, 1961; Moser et al., 1984; Matarese et al., 1989) and bars (Charter and Moser, 1996). The number of postanal bands and bars may or may not include the caudal or terminal notochord pigment. For clarity, in the descriptions of pigment patterns, the terms “band” and “bar” refer to aggregations of melanophores that approximate vertically oriented rectangles (including the caudal pigment). A band is always complete and a bar incomplete. A “stripe” approximates a line or elongate rectangle and is horizontally oriented. A “spot” is an approximately circular aggregation of melanophores. A “patch” is any other distinguishable aggregation of melanophores.

Larval morphology

A total of 115 eastern North Pacific Lepidopsetta larvae were measured with a calibrated digital image analysis system (larvae of the western North Pacific L. mochigarei were not available). This system consists of a video camera attached to a dissecting stereomicroscope or camera lens, a microcomputer with digital imaging board, and a video monitor. All measurements of larvae were taken from the left side. Unless otherwise noted, standard length (SL) is used throughout and is defined in our study as the length from snout tip to notochord tip (prior to development of the caudal fin) or to the posterior margin of hypural elements. Other measurements are defined as follows: head length (HL), snout tip to posterior edge of opercle (to pectoral-fin base in small larvae before opercular margin is visible); snout length (SNL), snout tip to anterior margin of orbit; orbit length (OL), greatest length of orbit; body depth (BD), vertical distance from dorsal to ventral body margin at a vertical line through center of anal opening; snout-to-anus length (SAL), distance along body midline from snout tip to a vertical line through center of anal opening. Standard proportional measurements were calculated (Table 2).

Table 2 Proportional measurements for Lepidopsetta larvae. SL = standard length; HL = head length. Statistical significance of raw data was evaluated by an ANCOVA analysis. See Table 10 for P-values. NS = not significant; S = significant.

Developmental stage L. polyxstra n. sp.

Yolksac

Sample size 6

Snout-to-anus length/SL 32.1 [+ or -] 1.7 (30.6-34.8)

Body depth/SL 3.8 [+ or -] 0.6 (2.7-4.3)

Head length/SL 11.6 [+ or -] 1.0 (10.1-12.6)

Snout length/HL 24.7 [+ or -] 2.4 (21.2-27.1)

Orbit length/HL 51.8 [+ or -] 5.6 (43.8-58.9)

Preflexion

Sample size 11

Snout-to-anus length/SL 33.5 [+ or -] 1.3 (32.3-36.0)

Body depth/SL 4.8 [+ or -] 0.7 (3.2-5.8)

Head length/SL 13.0 [+ or -] 1.3 (10.6-15.6)

Snout length/HL 20.5 [+ or -] 5.3 (13.3-29.7)

Orbit length/HL 48.1 [+ or -] 6.7 (34.1-61.9)

Flexion

Sample size 26

Snout-to-anus length/SL 35.5 [+ or -] 2.5 (31.3-40.0)

Body depth/SL 10.5 [+ or -] 4.1 (4.6-18.0)

Head length/SL 19.1 [+ or -] 3.6 (13.3-26.4)

Snout length/HL 20.5 [+ or -] 3.3 (12.8-26.2)

Orbit length/HL 31.1 [+ or -] 6.9 (20.6-44.9)

Postflexion

Sample size 10

Snout-to-anus length/SL 39.3 [+ or -] 3.3 (35.3-45.7)

Body depth/SL 28.5 [+ or -] 6.4 (20.0-38.6)

Head length/SL 26.7 [+ or -] 2.4 (22.4-30.6)

Snout length/HL 22.8 [+ or -] 2.1 (19.3-26.0)

Eye length/HL 20.9 [+ or -] 2.4 (17.5-24.0)

Total larvae measured 53

Developmental stage L. bilineata

Yolksac

Sample size 13

Snout-to-anus length/SL 32.9 [+ or -] 1.8 (30.1-36.8)

Body depth/SL 4.7 [+ or -] 0.8 (3.9-6.7)

Head length/SL 13.3 [+ or -] 1.4 (10.0-15.1)

Snout length/HL 22.6 [+ or -] 7.8 (13.1-36.5)

Orbit length/HL 51.9 [+ or -] 6.6 (36.7-61.5)

Preflexion

Sample size 13

Snout-to-anus length/SL 30.7 [+ or -] 2.3 (26.8-34.3)

Body depth/SL 5.0 [+ or -] 0.9 (3.3-6.7)

Head length/SL 12.7 [+ or -] 1.8 (10.4-15.8)

Snout length/HL 23.5 [+ or -] 6.3 (15.1-35.8)

Orbit length/HL 45.3 [+ or -] 5.6 (37.1-53.0)

Flexion

Sample size 17

Snout-to-anus length/SL 33.8 [+ or -] 2.1 (29.9-38.2)

Body depth/SL 11.1 [+ or -] 5.1 (5.5-21.9)

Head length/SL 17.9 [+ or -] 3.0 (14.6-24.4)

Snout length/HL 22.5 [+ or -] 4.7 (13.5-29.3)

Orbit length/HL 33.6 [+ or -] 4.1 (27.8-40.6)

Postflexion

Sample size 19

Snout-to-anus length/SL 34.6 [+ or -] 2.9 (29.3-38.2)

Body depth/SL 35.7 [+ or -] 3.0 (27.8-39.9)

Head length/SL 29.3 [+ or -] 1.7 (26.7-32.7)

Snout length/HL 20.7 [+ or -] 2.3 (16.8-24.2)

Eye length/HL 23.8 [+ or -] 3.2 (19.6-31.8)

Total larvae measured 62

Developmental stage Significance

Yolksac

Sample size

Snout-to-anus length/SL NS

Body depth/SL S

Head length/SL NS

Snout length/HL NS

Orbit length/HL NS

Preflexion

Sample size

Snout-to-anus length/SL S

Body depth/SL NS

Head length/SL NS

Snout length/HL NS

Orbit length/HL S

Flexion

Sample size

Snout-to-anus length/SL S

Body depth/SL S

Head length/SL NS

Snout length/HL NS

Orbit length/HL NS

Postflexion

Sample size

Snout-to-anus length/SL S

Body depth/SL S

Head length/SL S

Snout length/HL NS

Orbit length/HL S

Total larvae measured

Distributional analysis of larvae

The dataset used for mapping and calculation of mean density of larvae (number per 10 [m.sup.2]; Appendix Table 1) was a subset of the AFSC ichthyoplankton historical database (1972-1994) from the Bering Sea and Gulf of Alaska. This subset included all reidentified samples that had been originally identified as Psettichthys 2, Lepidopsetta 2, and L. bilineata and thus provided the only historic distributional records for larvae of L. polyxystra n. sp. Maps of material examined and distribution and mean density of species were generated by using the geographic information system Arc/Info and ArcView (Environmental Systems Research Institute, 1996). Tables were produced with ArcView and Systat software (SPSS, Inc., 1996). Densities were mapped by dividing the geographic area into contiguous square polygons, 625 [km.sup.2] each. Catches per unit of effort (CPUEs) from all tows (including 0 catches) within each square polygon were averaged, yielding a mean density. Polygons were shaded according to their mean density, or left unshaded when no tows occurred within a polygon.

Adult statistical analyses

The software programs S-Plus (Statistical Sciences, 1993) or Statgraphics Plus 2.1 (Manugistics, 1997) were used in statistical analyses performed on juveniles and adults. Unless otherwise indicated, tests were considered significant at P [is less than] 0.05, as adjusted by the Bonferroni correction (Sokal and Rohlf, 1995). For all characters, Bartlett’s test of homogeneity of variances (Sokal and Rohlf, 1995) was used to determine the appropriateness of an ANOVA to test for differences between species. When Bartlett’s test found significant differences in variances, pairwise comparisons were made. Pairs with significant differences in variances were then tested for differences in medians by using the nonparametric Kruskal-Wallis test (Sokal and Rohlf, 1995). The following meristic characters did not differ significantly in variance among species pairs and were subjected to an ANOVA: gill rakers of the upper and lower first arch and lower second arch; dorsal- and anal-fin rays; ocular-side and blind-side pectoral-fin rays; posterior and anterior ADB pores; interorbital scales; cheek scales; scales above, below, and total around the lateral line; ocular-side and blind-side suborbital pores; supraorbital pores.

For morphometric characters, significant differences between species were identified using an ANOVA of arcsine-transformed ratios of the measurement divided by SL or HL (Sokal and Rohlf, 1995) and an analysis of covariance (ANCOVA) with SL or HL as covariates when assumptions of homogeneity of slopes were satisfied. The following morphometric characters did not significantly differ in variances among species pairs and were subjected to an ANOVA: head length, snout length, ocular-side maxilla length, blind-side maxilla length, ocular-side mandible length, cheek length, interorbital width, dorsal orbit length, ocular-side pectoral-fin length, body depth, caudal peduncle depth, and caudal-fin length. Snout length, blind-side maxilla length, ocular-side mandible length, and interorbital width were also subjected to ANCOVA.

To aid in the discrimination and classification of species, standard principal components analysis (PCA) was conducted on all morphometric and meristic characters for adults of all species together and on the eastern North Pacific material separately. Raw morphometric data were log-transformed and the covariance matrix was subjected to PCA, as was the correlation matrix of raw meristics. Differences between species were illustrated by separately plotting principal components (PC) 2 and 3 of the morphometric analyses, PC1 and PC2 of the meristic analyses, and PC1 of the meristic analyses against PC2 of the morphometric analyses (Stauffer and Hert, 1992). For eastern North Pacific species, data points were also identified by reference to 11 geographic regions: Sea of Okhotsk, western Bering Sea, eastern Bering Sea, Aleutian Islands, Gulf of Alaska, British Columbia, Puget Sound, Washington coast, Oregon coast, California coast, and Baja California coast.

Larval statistical analyses

In an effort to identify additional characters to distinguish larvae of the two eastern North Pacific species, morphometric characters of yolksac preflexion, flexion, and postflexion larvae were further analyzed. Scatter plots for each measurement versus SL were made. Morphometric data were analyzed by the ANCOVA model for each parameter at each developmental stage and at all stages combined, which included species as a factor, standard length as a covariate, and a species/SL interaction (e.g. BD = C + Species + SL + (Species x SL)). A residual analysis was done for each model to determine the appropriateness of the model. Whenever the interaction was not significant (at the 5% level), a reduced model was used, and the interaction was dropped and the slopes were forced to be the same (BD = C + Species + SL). This procedure removed the effect of SL and allowed testing for significant differences between species. When the species were significantly different, the P-value was reported and graphs were generated to illustrate differences between elevations of the two regressions. Principal components analysis was also used to highlight differences.

Results

Adults

All three species differed significantly from each other in three meristic characters (Tables 3-7): posterior ADB pores (Table 5), total scales above and below the lateral line (Table 6), and ocular-side suborbital pores (Table 7). Lepidopsetta polyxystra n. sp. also differed from L. bilineata in number of gill rakers of the upper and lower first arch and lower second arch (Table 3), dorsal-fin rays (Table 4), ocular-side and blind-side pectoral-fin rays (Table 4), anterior ADB pores (Table 5), blind-side suborbital pores (Table 7), and supraorbital pores (Table 7), and differed from L. mochigarei in number of gill rakers of the upper and lower first arch and lower second arch (Table 3), blind-side pectoral-fin rays (Table 4), anterior and posterior ADB pores (Table 5), and cheek scales (Table 6).

Table 3 Counts of gill rakers for species of Lepidopsetta.

Upper first arch

Species 1 2 3 4 5

L. bilineata 12 167 104 4

L. mochigarei 1 18 8

L. polyxystra n. sp. 135 183 15

Upper first arch

Species 6 n x SD

L. bilineata 287 2.3 0.58

L. mochigarei 27 2.3 0.53

L. polyxystra n. sp. 3 336 3.7 0.61

Lower first arch

4 5 6 7 8

L. bilineata 5 105 160 17

L. mochigarei 1 5 18 3

L. polyxystra n. sp. 13 204 114

Lower first arch

9 n x SD

L. bilineata 287 5.7 0.62

L. mochigarei 27 5.9 0.66

L. polyxystra n. sp. 5 336 7.3 0.57

Lower second arch

5 6 7 8 9 10

L. bilineata 1 52 170 38 12 2

L. mochigarei 2 14 10 1

L. polyxystra n. sp. 4 17 125 139

Lower second arch

11 12 13 n x SD

L. bilineata 275 7.1 0.76

L. mochigarei 27 7.4 0.69

L. polyxystra n. sp. 40 10 1 336 9.7 0.92

Table 4 Fin-ray counts for species of Lepidopsetta.

Dorsal-fin rays

Species 64 67 68 69 70 71 72 73 74

L. bilineata 1 4 1 6 5 10

L. mochigarei 1 5

L. polyxystra n. sp. 1 3 5 6 12 9 13 12 13

Dorsal-fin rays

Species 75 76 77 78 79 80 81 82 83

L. bilineata 7 15 22 21 11 8 5 3 1

L. mochigarei 3 10 3 1 3 1

L. polyxystra n. sp. 8 20 14 10 14 7 4 1 2

Dorsal-fin rays

Species 84 85 86 89 n x SD

L. bilineata 1 1 4 1 127 77.14 3.57

L. mochigarei 27 76.52 2.46

L. polyxystra n. sp. 154 74.60 3.84

Anal-fin rays

Species 49 50 51 52 53 54 55

L. bilineata 7

L. mochigarei

L. polyxystra n. sp. 1 3 6 4 5 6 20

Anal-fin rays

Species 56 57 58 59 60 61 62

L. bilineata 2 13 26 17 29 15 9

L. mochigarei 1 3 5 4 5 6 1

L. polyxystra n. sp. 20 11 17 17 20 15 9

Anal-fin rays

Species 63 64 65 66 n x SD

L. bilineata 4 1 2 1 126 59.3 2.15

L. mochigarei 1 1 27 59.6 2.02

L. polyxystra n. sp. 3 2 153 57.5 3.28

Blind

pectoral-fin rays

Species 8 9 10 11 12

L. bilineata 10 52 62

L. mochigarei 2 17 6

L. polyxystra n. sp. 1 2 36 82 30

Blind

pectoral-fin rays

Species 13 n x SD

L. bilineata 2 126 11.4 0.66

L. mochigarei 2 27 11.3 0.72

L. polyxystra n. sp. 2 153 10.9 0.77

Ocular

pectoral-fin rays

Species 9 10 11 12 13

L. bilineata 5 12 69 32 5

L. mochigarei 5 14 7 1

L. polyxystra n. sp. 5 47 74 25 2

Ocular

pectoral-fin rays

Species n x SD

L. bilineata 123 11.2 0.81

L. mochigarei 27 11.1 0.77

L. polyxystra n. sp. 153 10.8 0.79

Table 5 Counts of lateral-line and supratemporal pores for species of Lepidopsetta.

Lateral-line pores

Species 70 71 72 73 74 75

L. bilineata 1 2 2

L. mochigarei

L. polyxystra n. sp.

Lateral-line pores

Species 76 77 78 79 80 81

L. bilineata 5 4 9 9 15 15

L. mochigarei

L. polyxystra n. sp. 2 2 1 5 6

Lateral-line pores

Species 82 83 84 85 86 87

L. bilineata 18 17 11 12 11 8

L. mochigarei

L. polyxystra n. sp. 3 12 12 24 15 14

Lateral-line pores

Species 88 89 90 91 92 93

L. bilineata 4 2 2 1

L. mochigarei

L. polyxystra n. sp. 16 10 7 10 5 6

Lateral-line pores

Species 94 95 96 97 98 99

L. bilineata

L. mochigarei 1 1 3

L. polyxystra n. sp. 2 4 1 1 1

Lateral-line pores

Species 100 101 102 103 104 105

L. bilineata

L. mochigarei 2 2 3 2 1 2

L. polyxystra n. sp. 1

Lateral-line pores

Species 106 107 108 109 115 117

L. bilineata

L. mochigarei 1 4 2 1 1

L. polyxystra n. sp.

Lateral-line pores

Species 119 n x SD

L. bilineata 148 82.1 3.67

L. mochigarei 1 27 104.4 5.84

L. polyxystra n. sp. 160 86.8 4.28

Posterior supratemporal pores

Species 6 8 9 10 11 12 13

L. bilineata 1 1 1 4 7

L. mochigarei 2 3 2 1 3 2 3

L. polyxystra n. sp. 3 2 2

Posterior supratemporal pores

Species 14 15 16 17 18 19

L. bilineata 6 13 5 15 14 12

L. mochigarei 3 3 1

L. polyxystra n. sp. 4 2 8 14 8 13

Posterior supratemporal pores

Species 20 21 22 23 24 25

L. bilineata 11 6 8 5 6 7

L. mochigarei 1 2

L. polyxystra n. sp. 16 16 6 11 10 8

Posterior supratemporal pores

Species 26 27 28 29 30 31

L. bilineata 4 1 2 4

L. mochigarei

L. polyxystra n. sp. 2 4 7 1 4 5

Posterior supratemporal pores

Species 33 34 35 n x SD

L. bilineata 1 134 19.1 4.73

L. mochigarei 26 12.6 4.28

L. polyxystra n. sp. 1 3 1 151 21.4 5.21

Anterior supratemporal pores

Species 2 3 4 5 6 7 8 9 10

L. bilineata 1 4 11 33 19 23 23 7 9

L. mochigarei 1 5 1 6 5 1 3 2

L. polyxystra n. sp. 1 1 9 15 30 25 30 25

Anterior supratemporal pores

Species 11 12 13 14 15 n x SD

L. bilineata 2 2 134 6.6 1.99

L. mochigarei 1 1 26 6.8 2.38

L. polyxystra n. sp. 8 3 1 2 1 151 8.3 1.99

Table 6 Counts of total scales above and below lateral line, scales above lateral line, scales below lateral line, and cheek scales for species of Lepidopsetta.

Total body scales

Species 65 66 67 68 69 70 71 72 73

L. bilineata 2 1 2 5 3 2 4 2 4

L. mochigarei

L. polyxystra n. sp. 1 1 1 1 3 1

Total body scales

Species 74 75 76 77 78 79 80 81 82

L. bilineata 7 5 3 9 6 3 7 4 4

L. mochigarei

L. polyxystra n. sp. 3 6 6 4 5 5 9 10 3

Total body scales

Species 83 84 85 86 87 88 89 90 91

L. bilineata 1 5 3 1 1 1

L. mochigarei 1

L. polyxystra n. sp. 7 7 3 1 3 1 5 2 2

Total body scales

Species 92 93 94 95 96 97 98 99

L. bilineata

L. mochigarei 1 1 1 5 2

L. polyxystra n. sp. 2 1 1

Total body scales

Species 100 101 102 103 104 105 106

L. bilineata

L. mochigarei 3 1 3 3 2 1 1

L. polyxystra n. sp.

Total body scales

Species 107 108 n x SD

L. bilineata 85 76.3 5.59

L. mochigarei 1 1 27 100.1 4.42

L. polyxystra n. sp. 94 80.8 5.95

Scales above lateral line

Species 25 26 27 28 29 30 31 32 33

L. bilineata 1 5 8 5 8 8 13 12 9

L. mochigarei

L. polyxystra n. sp. 5 5 5 11 9 16 17

Scales above lateral line

Species 34 35 36 37 38 39 40 41 42

L. bilineata 6 4 4 1 1

L. mochigarei 1 3 4 2 6 4 4

L. polyxystra n. sp. 7 9 6 2 1 1

Scales above lateral line

Species 43 44 45 n x SD

L. bilineata 1 85 31.0 3.13

L. mochigarei 1 1 1 27 39.9 2.49

L. polyxystra n. sp. 94 32.2 2.69

Scales below lateral line

Species 32 38 39 40 41 42 43 44 45

L. bilineata 1 1 1 3 10 5 6 5 9

L. mochigarei

L. polyxystra n. sp. 1 2 1 3 1 2 7

Scales below lateral line

Species 46 47 48 49 50 51 52 53 54

L. bilineata 11 10 7 6 3 3 2 2

L. mochigarei 1 1

L. polyxystra n. sp. 7 15 5 12 11 11 4 2 1

Scales below lateral line

Species 55 56 58 59 60 61 62 63 64

L. bilineata

L. mochigarei 2 3 4 2 4 3 4 1

L. polyxystra n. sp. 2 5 1 1

Scales below lateral line

Species 65 67 n x SD

L. bilineata 85 45.3 3.74

L. mochigarei 1 1 27 60.2 3.33

L. polyxystra n. sp. 94 48.6 3.95

Cheek scales

Species 7 8 9 10 11 12 13 14

L. bilineata 7 14 18 21 15 6

L. mochigarei 2 8 9 7

L. polyxystra n. sp. 3 5 14 36 29 10 2

Cheek scales

Species 15 16 n x SD

L. bilineata 2 1 84 11.6 1.55

L. mochigarei 1 27 11.9 1.11

L. polyxystra n. sp. 99 10.2 1.21

Table 7 Counts of ocular- and blind-side suborbital pores, preopercle pores, and supraorbital pores for species of Lepidopsetta.

Ocular-side suborbital pores

Species 13 14 15 16 17 18 19 20 21

L. bilineata 1 1 9 8 7 16

L. mochigarei

L. polyxystra n. sp. 1 2 2 4 2 7 11

Ocular-side suborbital pores

Species 22 23 24 26 27 28 29 30 31

L. bilineata 15 2 3 2 1 1

L. mochigarei 1 2 8 3 2

L. polyxystra n. sp. 11 17 10 6 3 3 3

Ocular-side suborbital pores

Species 32 33 34 35 36 37 38

L. bilineata

L. mochigarei 3 3 1 2 1

L. polyxystra n. sp.

Ocular-side

suborbital pores

Species n x SD

L. bilineata 72 21.2 2.71

L. mochigarei 27 30.7 3.51

L. polyxystra n. sp. 93 22.7 3.10

Blind-side suborbital pores

Species 6 7 8 9 10 11 12 13 14

L. bilineata 1 2 12 24 21 9 3 2

L. mochigarei 2 3 7 8 4

L. polyxystra n. sp. 5 17 32 31 3 4 1

Blind-side suborbital pores

Species 15 16 17 19 20 n x SD

L. bilineata 74 9.5 1.32

L. mochigarei 2 1 27 12.9 1.94

L. polyxystra n. sp. 93 10.3 1.16

Supraorbital pores

Species 1 2 3 4 5 6 7

L. bilineata 2 6 20 70 91 51

L. mochigarei 17 1 1

L. polyxystra n. sp. 118 105 29 7 2 3 2

Supraorbital pores

Species 8 9 n x SD

L. bilineata 10 5 255 5.8 1.19

L. mochigarei 19 1.2 0.50

L. polyxystra n. sp. 266 1.8 1.03

Preopercle pores

Species 5 6 7 8 9 10 11

L. bilineata 2 60 9

L. mochigarei 1 4 2 8 6

L. polyxystra n. sp. 1 32 53 3

Preopercle pores

Species 12 13 n x SD

L. bilineata 71 11.6 1.55

L. mochigarei 1 2 24 11.9 1.11

L. polyxystra n. sp. 89 10.2 1.21

All three species were found to differ significantly in the means of two morphometric characters with either ANOVA or ANCOVA: interorbital width and ocular-side mandible length (Table 8). Lepidopsetta polyxystra n. sp. also differed from L. bilineata in seven additional characters: head length, blind-side maxilla length, cheek length, body depth, ocular-side pectoral-fin length, caudal-fin length, and caudal peduncle depth; and differed in medians in ocular-side maxilla length, blind-side pectoral-fin length, and body depth. Lepidopsetta polyxystra n. sp. also differed from L. mochigarei in means of two characters: dorsal orbit length and cheek length; and in medians of head length, body depth, depth at caudal base, and caudal length. Lepidopsetta bilineata also differed from L. mochigarei in means of two additional characters: blind-side maxilla length and dorsal orbit length; and differed in medians in head length, ocular-side maxilla length, cheek length, body depth, caudal peduncle depth, and depth at caudal base. No differences in means were found in snout length or ocular-side maxilla length.

Table 8 Proportional morphometrics of adults of Lepidopsetta. HL = head length; SL = standard length. Data are in hundreds of SL or HL with mean [+ or -] standard deviation (range). Statistical significance was evaluated by ANOVA and ANCOVA, when appropriate, at 0.05 level. All = Differences between all three species were significant; bp = Differences between L. bilineata and L. polyxystra n. sp. were significant; pm = Differences between L. polyxystra n. sp. and L. mochigarei were significant; ns = No significant differences were found.

Species

L. bilineata

Sample size 83

Head length/SL 28.20 [+ or -] 1.64 (24.01-32.86)

Snout length/HL 16.09 [+ or -] 2.07 (12.37-22.45)

Ocular-side maxilla

length/HL 27.88 [+ or -] 1.25 (25.26-32.53)

Ocular-side mandible

length/HL 41.91 [+ or -] 1.69 (38.33-46.10)

Interorbital width/HL 3.54 [+ or -] 0.79 (2.14-6.02)

Dorsal orbit length/HL 28.06 [+ or -] 1.91 (23.38-32.46)

Ventral orbit length/HL 3.54 [+ or -] 0.79 (2.14-6.02)

Cheek length/HL 34.17 [+ or -] 2.84 (28.06-41.84)

Body depth/SL 46.98 [+ or -] 2.84 (40.80-57.04)

Ocular-side pectoral

length/SL 15.19 [+ or -] 1.34 (12.20-18.20)

Caudal-fin length/SL 20.79 [+ or -] 1.57 (17.62-23.63)

Caudal peduncle

depth/SL 10.20 [+ or -] 0.80 (8.18-12.29)

Caudal peduncle

length/SL 9.11 [+ or -] 0.72 (7.18-10.87)

Ocular-side pelvic

length/SL 9.83 [+ or -] 0.98 (7.64-12.50)

Sample size 77

Blind-side maxilla

length 31.26 [+ or -] 2.25 (25.16-41.18)

Sample size 78

Blind-side pectoral

length/SL 9.91 [+ or -] 0.77 (7.33-12.22)

Sample size 67

Cheek depth/HL 19.15 [+ or -] 2.13 (13.92-24.19)

Caudal depth at

hypurals/SL 12.13 [+ or -] 0.69 (10.46-14.25)

Sample size 8

Lateral line length/

lateral line depth 30.20 [+ or -] 3.78 (24.84-37.06)

Species

L. mochigarei

Sample size 26

Head length/SL 26.23 [+ or -] 1.71 (23.23-30.54)

Snout length/HL 16.13 [+ or -] 1.69 (12.72-20.21)

Ocular-side maxilla

length/HL 27.21 [+ or -] 1.41 (24.63-29.41)

Ocular-side mandible

length/HL 39.61 [+ or -] 1.83 (36.57-44.64)

Interorbital width/HL 2.68 [+ or -] 0.69 (1.32-4.17)

Dorsal orbit length/HL 30.08 [+ or -] 1.83 (26.09-34.74)

Ventral orbit length/HL 2.68 [+ or -] 0.69 (1.32-4.17)

Cheek length/HL 31.09 [+ or -] 1.71 (28.57-34.98)

Body depth/SL 51.79 [+ or -] 3.23 (45.54-61.04)

Ocular-side pectoral

length/SL 14.58 [+ or -] 1.24 (12.60-17.42)

Caudal-fin length/SL 21.22 [+ or -] 1.69 (18.77-25.14)

Caudal peduncle

depth/SL 11.03 [+ or -] 0.94 (9.71-13.96)

Caudal peduncle

length/SL 8.89 [+ or -] 0.54 (7.77-10.28)

Ocular-side pelvic

length/SL 9.82 [+ or -] 0.94 (7.40-11.75)

Sample size 26

Blind-side maxilla

length 29.73 [+ or -] 1.56 (27.30-32.87)

Sample size 26

Blind-side pectoral

length/SL 9.58 [+ or -] 0.90 (7.89-11.69)

Sample size 26

Cheek depth/HL 19.64 [+ or -] 2.60 (14.83-25.53)

Caudal depth at

hypurals/SL 13.12 [+ or -] 0.70 (11.66-14.61)

Sample size 6

Lateral line length/

lateral line depth 33.42 [+ or -] 3.78 (27.86-38.98)

Species

L. polyxystra

Sample size 94

Head length/SL 27.24 [+ or -] 1.62 (21.75-31.54)

Snout length/HL 16.02 [+ or -] 1.84 (12.06-20.12)

Ocular-side maxilla

length/HL 26.30 [+ or -] 1.89 (20.42-31.47)

Ocular-side mandible

length/HL 40.73 [+ or -] 2.20 (34.39-50.00)

Interorbital width/HL 4.35 [+ or -] 0.80 (2.49-6.56)

Dorsal orbit length/HL 28.13 [+ or -] 1.95 (22.18-32.86)

Ventral orbit length/HL 4.35 [+ or -] 0.80 (2.49-6.56)

Cheek length/HL 32.54 [+ or -] 2.77 (26.67-38.52)

Body depth/SL 49.02 [+ or -] 3.44 (41.17-58.93)

Ocular-side pectoral

length/SL 14.66 [+ or -] 1.52 (10.73-18.23)

Caudal-fin length/SL 22.72 [+ or -] 1.77 (17.78-29.13)

Caudal peduncle

depth/SL 10.91 [+ or -] 0.71 (9.36-13.13)

Caudal peduncle

length/SL 8.90 [+ or -] 0.62 (7.01-10.12)

Ocular-side pelvic

length/SL 9.95 [+ or -] 0.97 (7.84-12.15)

Sample size 93

Blind-side maxilla

length 28.70 [+ or -] 2.01 (22.22-35.66)

Sample size 93

Blind-side pectoral

length/SL 10.16 [+ or -] 1.17 (6.58-12.66)

Sample size 92

Cheek depth/HL 18.62 [+ or -] 2.17 (13.56-22.73)

Caudal depth at

hypurals/SL 12.35 [+ or -] 0.88 (10.06-16.91)

Sample size 15

Lateral line length/

lateral line depth 31.72 [+ or -] 3.41 (26.29-38.20)

Significance

ANOVA ANCOVA

Sample size

Head length/SL bp

Snout length/HL ns ns

Ocular-side maxilla

length/HL ns

Ocular-side mandible

length/HL bm All

Interorbital width/HL All All

Dorsal orbit length/HL pm/bm

Ventral orbit length/HL

Cheek length/HL bp/pm

Body depth/SL bp

Ocular-side pectoral

length/SL bp

Caudal-fin length/SL bp

Caudal peduncle

depth/SL bp

Caudal peduncle

length/SL

Ocular-side pelvic

length/SL

Sample size

Blind-side maxilla

length bp/bm bp/bm

Sample size

Blind-side pectoral

length/SL

Sample size

Cheek depth/HL

Caudal depth at

hypurals/SL

Sample size

Lateral line length/

lateral line depth

In the standard PCA of all three species together, loadings of morphometric PC1 were all positive and thus exhibited a strong size effect (Table 9), accounting for 95% of morphometric variation. Principal components 2 and 3 accounted for 56% of the remaining morphometric variation. Principal component 2 loadings described a gradient based on interorbital width and dorsal orbit length; PC3 loadings, a gradient on caudal peduncle depth, cheek depth, caudal depth at hypural margins, and body depth. The plot of PC2 versus PC3 revealed extensive overlap in morphometric characters (Fig. 2A). Principal components 1 and 2 of the meristics analysis accounted for 46% of the meristic variance: PC1 strongly loaded on lateral line pores and scales above and below the lateral line, preopercle pores, and suborbital pores, whereas PC2 strongly loaded on gill rakers of the lower first and second arches and upper first arch and supraorbital pores (Table 10). By plotting PC1 of the meristic analysis and PC2 of the morphometric analysis (Fig. 2C), we found a distinct cluster representing L. mochigarei and an overlap of clusters representing L. bilineata and L. polyxystra n. sp.

[Figure 2 ILLUSTRATION OMITTED]

Table 9 Character loadings for principal component analysis of morphometric characters of adult Lepidopsetta bilineata, L. mochigarei, and L. polyxystra n. sp.

Loadings

Characters PC 1 PC2 PC3

Standard length 0.22484 0.06236 0.15282

Head length 0.22819 0.03318 -0.06512

Snout length 0.21789 0.00863 -0.04715

Ocular-side maxilla length 0.23138 0.05417 -0.30392

Blind-side maxilla length 0.23849 0.0835 -0.30268

Ocular-side mandible length 0.2334 0.03438 -0.20318

Dorsal orbit length 0.21415 0.11385 -0.09032

Ventral eye length 0.20185 0.09275 -0.20682

Interorbital width 0.22077 -0.96363 -0.03502

Cheek length 0.24796 0.00482 -0.13585

Cheek depth 0.25623 0.08324 -0.39732

Body depth 0.23716 0.08671 0.33759

Ocular-side pectoral length 0.24489 0.0723 -0.01453

Blind-side pectoral length 0.24277 0.01015 0.184878

Ocular-side pelvic length 0.22725 0.0275 0.076978

Caudal peduncle length 0.22574 0.08882 0.121448

Caudal peduncle depth 0.22183 0.00716 0.401018

Caudal depth at hypurals 0.22707 0.07164 0.377584

Caudal-fin length 0.20981 -0.00818 0.20981

Table 10 Character loadings for principal component analysis of meristic characters for Lepidopsetta bilineata, L. mochigarei, and L. polyxystra n. sp. ADB = supratemporal branch of the lateral line.

Loadings

Characters PC1 PC2

Upper 1st arch gill rakers 0.0722 0.4021

Lower 1st arch gill rakers 0.0646 0.4414

Upper 2nd arch gill rakers 0.0067 0.1393

Lower 2nd arch gill rakers 0.0181 0.406

Dorsal-fin rays -0.1314 -0.1926

Anal-fin rays -0.1265 -0.1939

Ocular-side pectoral-fin rays -0.0975 -0.1331

Blind-side pectoral-fin rays -0.0759 -0.1527

Lateral-line pores -0.4017 0.1027

ADB1 pores 0.2092 0.1578

ADB2 pores 0.0079 0.1962

Interorbital scales -0.0508 0.2163

Cheek scales -0.1907 -0.2532

Scales above lateral line -0.384 0.0291

Scales below lateral line -0.392 0.0873

Ocular-side suborbital pores -0.3635 0.0177

Blind-side suborbital pores -0.3007 0.0961

Preopercle pores -0.3643 0.0394

Supraorbital pores 0.203 -0.3731

In the PCA of only the eastern North Pacific species, loadings of morphometric PC1 were all positive and thus exhibited a strong size effect, accounting for 96% of morphometric variation. Principal components 2 and 3 accounted for 33% and 19% of the remaining morphometric variation, respectively. Principal component 2 loadings described a gradient based primarily on interorbital width, as well as on ventral orbit length and blind-side maxilla length; PC3 loadings described a gradient on interorbital width, caudal peduncle depth, and body depth. The plot of PC2 versus PC3 revealed extensive overlap in morphometric characters (Fig. 3A). Principal components 1 and 2 of the meristics analysis accounted for 28% and 17%, respectively, of the meristic variance: PC1 strongly loaded on gill rakers and supraorbital pores and PC2 strongly loaded on dorsal-fin rays, anal-fin rays, and cheek scales. The plot of meristic PC1 versus PC2 revealed two distinct clusters (Fig. 3B). By plotting PC1 of the meristic analysis and PC2 of the morphometric analysis (Fig. 3C), we found two distinct clusters representing L. bilineata and L. polyxystra n. sp.

[Figure 3 ILLUSTRATION OMITTED]

When data points of eastern North Pacific species were identified by geographic regions, no significant clustering was apparent within groups of either L. bilineata or L. polyxystra n. sp. Extensive overlap was found between clusters representing each of these general geographic areas.

Larvae

Raw morphometric data were subjected to an ANCOVA analysis, which indicated differences in several morphological characters (snout-to-anus length, body depth, and orbit length) between larvae of L. bilineata and L. polyxystra n. sp. at various stages of development (Table 11). Several characters showed significant differences in postflexion larvae (P [is less than] 0.0001), including snout-to-anus length, head length, and orbit length. Snout-to-anus length is shorter in larvae of L. bilineata during the preflexion, flexion, and postflexion stages (Fig. 4). Larvae of L. bilineata are deeper-bodied during the yolksac, preflexion, and flexion stages of development (Fig. 5). The PCA plots of PC2 and PC3 for each developmental stage produced broadly overlapping clusters.

[Figure 4-5 ILLUSTRATION OMITTED]

Table 11 P-values for ANCOVA of larvae of Lepidopsetta bilineata and L. polyxystra n. sp. Bold values indicate that the species are different from each other at the 0.05 significance level.

Stage of development

Character Yolksac Preflexion Flexion Postflexion

Snout-to-anus

length NS P=0.001 P=0.035 P<0.001

Body depth P=0.001 NS P=0.037 P<0.001

Head length NS NS NS P=0.001

Snout length NS NS NS NS

Orbit length NS P=0.020 NS P<0.001

Systematics

Genus Lepidopsetta Gill, 1862

Lepidopsetta Gill, 1862:330 (type species Platichthys umbrosus Girard, 1856 [originally cited as Psettichthys umbrosus but subsequently corrected by Gill (1864)], by monotypy).

Diagnosis

Pleuronectid genus Lepidopsetta is distinguished by the following combination of characters in adults: lateral line with a high arch, 6-8 scale rows between highest point of arch and base of arch over the pectoral fin; supratemporal branch of lateral line with a moderately elongate posterior extension of up to 35 pored scales; mouth small, maxilla extending to anterior quarter of ventral orbit; expanded anteriormost anal pterygiophore (“anal spine”) present, often projecting beyond body wall in damaged specimens; supraorbital pores present; scales on ocular side often tuberculate, especially on the head and anterior portion of body; scales on blind side always cycloid anteriorly.

Larvae are distinguished from other pleuronectid genera by the following combination of characters: dorsal and anal finfold pigment present, 1-3 postanal bars (2-4 distinctive pigment regions along dorsal midline), a series of ventral midline melanophores, absence of slashlike hypaxial pigment along the postanal body, few to many pigment spots around the notochord tip, and 37-44 total myomeres (Table 12).

Table 12 Summary of selected larval characters helpful in distinguishing eastern North Pacific pleuronectid larvae during preflexion and early flexion stages (Matarese et al., 1989, in part). Characters are presented for taxa where at least some early life-history stages are known. Only general trends are presented because pigment may vary among specimens. When specimens were not available, subjective decisions were based on previously published material. PVM = postventral melanophores.

Pigment characters

Postanal Notochord

Genus bars Hypaxial PVM tip

Embassichthys 3 Absent Absent Present

Eopsetta 3 Absent Absent Present

Glyptocephalus 4 Absent Absent Present

Hippoglossoides 3-4 Present Absent Present

Isopsetta 3 Absent Absent Present

Lepidopsetta 1-3 Absent Present Present

Microstomus 3-4 Absent Absent Present

Psettichtys 1 Absent Absent Present

Acanthopsetta Absent Present Present Present

Atheresethes Absent Absent Absent Present/

absent

Hippoglossus Absent Present Absent Present

Limanda Absent Present Present Absent

Liopsetta Absent Absent Absent Present

Lyopsetta Absent Present Absent Present

Parophrys Absent Present Present Present

Platichthys Absent Absent Present Present

Pleuronectes Absent Absent Present Absent

Pleuronichthys Absent Absent Absent Present

Reinhardtius Absent Present Absent Present

Pigment characters

Genus Dorsal Anal Head Total

finfold finfold spines myomeres

Embassichthys Present Present Absent 57-65

Eopsetta Absent Absent Preopercular 41-45

Glyptocephalus Present/ Present/ Preopercular 52-66

absent absent

Hippoglossoides Present Present Absent 44-51

Isopsetta Absent Present Absent 41-42

Lepidopsetta Present Present Absent 37-44

Microstomus Present Present Otic 50-55

Psettichtys Present Present Absent 38-41

Acanthopsetta Absent Absent Absent 39-40

Atheresethes Absent Absent Preopercular/ 47-50

supraocular crest

Hippoglossus Present Present Absent 49-51

Limanda Absent Present Absent 40-41

Liopsetta Absent Absent Absent 37-41

Lyopsetta Present Present Absent 43-47

Parophrys Absent Present Absent 42-47

Platichthys Present Present Absent 35-38

Pleuronectes Absent Present Absent 41-42

Pleuronichthys Present Present Absent/otic 38-41

Reinhardtius Present Present Absent 61-64

Description of adults

Body ovate, greatest depth 31.7-61.0% SL, scales above lateral line 25-45, scales below lateral line 27-67; head acute to rounded, length 21.8-32.9% SL; dorsal margin of head concave to nearly linear, snout length 3.1-6.2% SL (12.1-22.4% HL); mouth small, premaxilla with fleshy lips and prominent, broad-based posterior lobe, maxilla extending to anterior rim of ventral orbit, ocular-side maxilla shorter than blind-side maxilla, ocular-side maxilla length 20.4-32.5% HL, blind-side maxilla length 22.2-41.2% HL; ocular-side mandible length 34.4-50.0% HL, about equal in length to blind-side mandible; teeth conical and slightly lobate or truncate, largest are anterior becoming gradually smaller posteriorly, fewer on ocular-side premaxilla and dentary, 4-10 on ocular-side premaxilla, 20-27 on blind-side premaxilla, 8-15 on ocular-side dentary, 20-29 on blind-side dentary; gill rakers of first arch moderately slender to broad and robust, 6-14 total, 1-6 on upper and 4-9 on lower arch; gill rakers of second arch broad and robust, 6-14 total, 0-4 on upper and 5-13 on lower arch; dorsal orbit round (nearly equal to eye length) to elliptical (posterior rim elongate and much greater than eye length), orbit lenth 22.2-34.7% HL, dorsal eye length 17.9-32.9% HL; ventral eye approximately equal in length to ventral orbit, length 17.2-35.4% HL; interorbital with up to 5 scales (often lost in preserved material), width 1.3-6.6% HL; cheek with 7-16 scales, length 26.7-65.9% HL, depth 13.6-33.6% HL; preopercular pores 5-13; ocular-side suborbital pores 13-38; blind-side suborbital pores 6-20; lateral line with a high arch, 6-8 scale rows high over the pectoral fin, arch length about three times its depth, length 26.1-69.9% HL, lateral line pores 70-119; anterior supratemporal short, pores 2-15; posterior supratemporal branch moderately elongate, pores 6-35; supraorbital canal short or long, pores 1-9; pectoral fins with 8-13 rays, dorsal 2-3, and ventralmost simple and all others branched, ocular-side pectoral fin longer than blindside pectoral fin, length 10.7-18.2% SL (38.3-70.4% HL), blind-side pectoral fin about equal in length to pelvic fins, length 6.6-12.7% SL (23.2-49.6% HL); pelvic fins with 6 branched rays, ocular-side pelvic-fin length about equal to blind-side pelvic-fin length, 7.4-12.5% SL (26.7-45.9% HL); dorsal-fin origin over anterior portion of dorsal orbit, dorsal fin with 64-89 simple soft rays, smallest rays anterior and posterior, longest rays at midbody, height 10.2-16.6% SL; dorsal-fin pterygiophores 68-78 in specimens with 64-83 dorsal-fin rays, 8-11 anterior to first neural spine, the first bifurcate and supporting two rays; anal fin with 49-77 simple soft rays in specimens with 54-63 anal-fin rays supported by 53-63 pterygiophores, fin about equal in height to dorsal fin; anal-fin rays 1 and 2 supported by greatly expanded anterior pterygiophore, sharp point of which often projects through skin between anal-fin origin and anus; caudal peduncle length 7.0-10.9% SL (25.5-42.2% HL); least caudal peduncle depth 8.2-14.0% SL (29.6-50.3% HL; 90.0-172.0% caudal peduncle length); greatest caudal peduncle depth 10.1-16.9% SL (37.2-57.7% HL); caudal fin truncate to rounded, with 18-19 rays, total of 8-10 both dorsally and ventrally, five on each of complex hypurals 1+2 and 3+4, 3 on hypural 5, one on the epural or preural 2 neural spine, and four on the parhypural, length 15.3-29.1% SL; vertebrae 39-42, 11 precaudal and 28-31 caudal, neural spines 1-3 and haemal spines 1-4 anteroposteriorly expanded, epineurals present on vertebrae 2-11, ribs present on vertebrae 3-11. Meristics are summarized in Tables 3-7.

Ocular-side scales ctenoid, often tuberculate, especially on anterior portion of the body around the head and pectoral girdle; tubercules columnar, narrow from base to tip, up to 10 tubercules on scales of head, fewer on tuberculate body scales; blind-side scales nearly always cycloid except in ambicolored specimens, never tuberculate, posterior scales occasionally with central cteni along lateral midline; broad, flaplike urogenital pore on ocular side, dorsal to anus; anus just anterior to anal-fin origin; color of ocular side varies with substrate, in life olivaceous greenish brown with various blotches of dark brown and spots of light areas scattered over body, brown streaks in dorsal, anal, and caudal fins, often with a series of four to five large light spots at base of dorsal fin and four similar spots at base of anal fin; in preservation ocular side yellowish brown with blotches; blind side bright white to cream in life, cream to yellowish in preservation. Moderate sized pleuronectids, reaching a standard length of 600 mm (Hart, 1973).

Description of larvae

Snout-to-anus length remaining constant during development, length 32.1-39.3% SL; preflexion body slender, body depth increasing with development, sharply after flexion, from a depth of 3.8 to 35.7% SL; finfolds of moderate size; head length increasing with development, sharply after flexion, from a length of 11.6 to 29.3% SL; snout length remaining constant during development, length 20.5-24.7% HL; orbit length decreasing with development, from 51.9 to 20.9% HL.

Size at stage of development variable among species (Table 13). Larvae hatching at less than 3.0-4.45 mm; yolk absorbed by 2.7-4.5 mm. Prefiexion larvae ranging in size from 3.8 to 6.6 mm; flexion larvae between 6.4 and 11.1 mm; postflexion larvae between 10.8 and 18.6 mm. Transformation occurring at sizes as small as 10.0 mm; juvenile stage usually attained by 35.0 mm (Table 13).

Table 13 Length at stage of development for species of Lepidopsetta (mm SL). For a more complete explanation of each developmental stage, see Kendall et al. (1984).

Stage L. bilineata L. mochigarei

Egg ~1.0 0.90

Hatch <3.0 3.95-4.48

Yolk absorption 2.7-4.5 unknown

Preflexion 3.8-6.6 unknown

Flexion 6.4-11.1 ~8.9-10.6

Postflexion 10.8-16.0 unknown

Transformation as small as 10.0 as small as 15.3

Early juvenile complete as small unknown

as 13.0 (13.0-33.9)

Postsettlement juvenile <35.0 unknown

Stage L. polyxystra n. sp.

Egg ~1.0

Hatch >3.0

Yolk absorption 3.3-4.2

Preflexion 4.2-6.2

Flexion 6.2-12.6

Postflexion 12.2-18.6

Transformation as small as 15.0

Early juvenile complete as small as

26.0

Postsettlement juvenile >35.0

Preanal pigment present initially along lower jaw, increasing posteriorly with development. Pigment occurring ventrally along gut and dorsally on anus; by flexion a distinct patch of melanophores occurring along the posterior edge of the gut, with pigment increasing laterally with development. Pigment may be present or absent on pectoral-fin rays and base.

Postanal pigment in preflexion larvae may be present as melanophores along distal edges of dorsal and anal finfolds, but typically with isolated patches along finfolds and pigment spots or patches along distal edges of anal finfold; when present, two to four distinct pigment regions along dorsal midline; a series of melanophores present from the gut along the ventral midline posterior to anus to just posterior to ventral stripe of posteriormost bar; a few melanophores on the caudal peduncle and above and below the notochord tip.

Distribution

The genus is endemic to the North Pacific and is widespread on the continental shelf (Figs. 6-13). Its range extends from Yongil Bay, Korea, in the southern Sea of Japan and along the northern coasts of Japan, north through the Sea of Okhotsk and the Bering Sea, the most northerly record being from the Gulf of Anadyr and the vicinity of St. Lawrence Island. It is recorded from the Aleutian Islands and into the eastern North Pacific throughout the Gulf of Alaska, south into Puget Sound and along the west coast of Washington, Oregon, California, and Mexico. Its most southerly record in the eastern Pacific is from the Cortez Banks of Baja California, Mexico.

[Figures 6-13 ILLUSTRATION OMITTED]

Larvae of the eastern North Pacific species range from the Bering Sea and Aleutian Islands along the Pacific coast to Baja California (Figs. 12-13). Both species co-occur in the eastern Aleutian Islands to the Washington coast; only larvae of L. polyxystra n. sp. have been routinely collected from the Bering Sea and only larvae of L. bilineata have been collected from California and south in California Cooperative Oceanic Fisheries Investigations (CalCOFI) samples. A subset of larvae collected by AFSC surveys in the Gulf of Alaska and Bering Sea was examined to compare differences in temporal distribution and mean density (number/10 [m.sup.2]; Appendix Table 1). During the 22-year sampling period, larvae of L. polyxystra n. sp. appear first in our March collections (Table 14), whereas larvae of L. bilineata first appear in April. The largest catches of L. polyxystra n. sp. occurred in May, whereas the largest catches of L. bilineata occurred in June. Overall, in the subset of cruises from 1972 to 1994 considered in our study, L. polyxystra n. sp. were more abundant and mean density was higher (5.34/10 [m.sup.2]) than that of L. bilineata (0.414/10 [m.sup.2]).

Table 14 Monthly collections of larvae of Lepidopsetta in the Gulf of Alaska and Bering Sea (1972-1994). Mean density (number per 10 [m.sup.2]) followed below by the standard deviation of the mean in parentheses.

Month

Taxon Jan Feb Mar Apr May

L. bilineata 0 0 0 0.0139 0.568

(0) (0) (0) (0.0059) (0.0592)

L. polyxystra n. sp. 0 0 0.456 5.80 7.06

(0) (0) (0.204) (0.633) (0.383)

Total observations 115 163 704 2926 2678

Month

Taxon Jun Jul Aug Sep

L. bilineata 1.73 0.15 0 0.116

(0.243) (0.0553) (0) (0.0872)

L. polyxystra n. sp. 4.44 0.659 0 0.09

(0.481) (0.018) (0) (0.0634)

Total observations 861 333 0 144

Month

Taxon Oct Nov Dec Overall

L. bilineata 0.0311 0 0 0.414

(0.0311) (0) (0) (0.0357)

L. polyxystra n. sp. 0.0386 0 0 5.34

(0.0386) (0) (0) (0.289)

Total observations 138 585 11 7502

Habitat

Species of the continental shelf were collected over sand and gravel, commonly at depths of 200 m and less, to as deep as 575 m (Allen and Smith, 1988). For eastern North Pacific species, highest densities of larvae were found over depths of less than 500 m, although they were also collected over deeper water.

Life history

Described eggs of Lepidopsetta are demersal, off-round, and have a sticky chorion, causing them to adhere to each other or to a substrate, and range in size from 0.86 to 1.08 mm in diameter (Yusa, 1958; Pertseva-Ostroumova, 1961; Penttila, 1995). The reproductive season extends from winter to early summer, generally earlier in southern species, and spawning occurs at depths of less than 220 m. All sexually mature Lepidopsetta apparently migrate from shallow shelf areas in the fall to deeper upper slope waters during winter and migrate back into shallower shelf waters during spring and summer. Immature Lepidopsetta remain in shallow waters throughout the winter and migrate into shallower coastal waters in the spring and summer.

Garrison and Miller (1982) provided a summary of reproductive characteristics of Lepidopsetta from the western and eastern Pacific Ocean, which included L. bilineata and L. polyxystra n. sp. Blackburn (1973) described the ichthyoplankton from Skagit Bay, Washington, located in the northernmost region of Puget Sound and included descriptions and illustrations of larvae of Lepidopsetta. The Washington Department of Fisheries (WDF) has reported eggs of Lepidopsetta (L. bilineata or L. polyxystra n. sp., or both) from late December through early March in sandy gravel of upper intertidal beaches in several sites in central and southern Puget Sound (Penttila, 1995). These eggs were described as demersal and adhesive (“clinging to upper beach surface material,” p. 238). Eggs were collected at the same times and location as those of surf smelt (Hypomesus pretiosus) and Pacific sand lance (Ammodytes hexapterus).

Etymology

The name Lepidopsetta is derived from the Greek lepido meaning “scaled,” a probable reference to the strongly ctenoid scales found on the ocular side of most individuals, and psetta, meaning “flatfish”.

Comments

Sakamoto (1984a) synonymized Lepidopsetta (as well as five other pleuronectid genera: Limanda, Parophrys, Isopsetta, Pseudopleuronectes, and Liopsetta) with Pleuronectes, based on a phenetic analysis of detailed morphological data. Several recent authors have criticized Sakamoto’s (1984a) phylogenetic techniques (Chapleau [1993]; Rass [1996]; Berendzen, [1998]; Cooper and Chapleau [1998]) and have refuted the monophyly of Pleuronectes sensu Sakamoto (Rass [1996]; Cooper and Chapleau [1998]). Cooper and Chapleau (1998) recently conducted a cladistic analysis and resurrected these genera from synonymy with Pleuronectes. We follow the consensus among most current scientists (Nelson, 1994; Cooper and Chapleau, 1996, 1998; Rass, 1996) and recognize each of these genera as distinct.(4)

Supraorbital pores have not been reported in other flat-fishes. The supraorbital canal is an extension of a nerve branch arising from between the frontals at the posterior rim of the dorsal orbit. In Lepidopsetta the pores of the canal are supported by a series of 1-6 bones (Fig. 14), similar in size and shape to the suborbitals. Typically each bone shares in the support of one or two pores.

[Figure 14 ILLUSTRATION OMITTED]

Among other North Pacific pleuronectids examined, Parophrys vetulus(5) has 6-11 pores and Isopsetta isolepis 2-12 pores, whereas Limanda aspera and L. pinnifasciatus, Psettichthys melanostictus, and Inopsetta ischyra (a possible hybrid of Platichthys stellatus and Parophrys vetulus) have a single pore. In Hippoglossoides elassodon and H. robustus, a series of pores is found at the anterior extension of the supratemporal line, as depicted by Lindberg and Fedorov (1993) for H. elassodon (their Fig. 33). These pores are apparently part of the anterior extension of the trunk lateral line, but ventrally the pores extend near the dorsal orbital rim. In one specimen of H. elassodon examined (UW 040271), the pores extended nearly the same length along the rim of the dorsal orbit as the length for the same pores in some L. bilineata.

Pleuronectid larvae are difficult to diagnose on the basis of a simple set of characters. Pleuronectids are oviparous and spawn planktonic or demersal eggs (about 0.66-4.5 mm), with homogenous yolk that is either pigmented or not, and usually contains no oil. Among eastern North Pacific pleuronectids, Lepidopsetta is the only genus that produces demersal eggs, which are off-round and have a sticky chorion that causes them to adhere to each other or to a substrate (Pertseva-Ostroumova, 1961; Penttila, 1995). Pleuronectid larvae hatch between 1.7 and 16.0 mm; yolksac and preflexion larvae are slender, becoming deeper-bodied during the postflexion stage. Larvae of Lepidopsetta hatch at [is less than] 3.0-4.5 mm; preflexion larvae are slender and have moderate-size finfolds.

Larval pigmentation (including finfold) varies considerably in pleuronectids. A combination of pigment pattern (postanal bands, bars, and finfold), meristic characters, and size at development is needed for identification (Matarese et al., 1989; Charter and Moser, 1996). Larvae with special characters (e.g. Atheresthes larvae with head spines) or high myomere counts (e.g. Embassichthys) are easily distinguished. Other pleuronectid larvae are usually distinguished by a combination of pigmentation characters including the number of postanal bands and bars and finfold pattern (Table 12).

No single early life history character distinguishes larvae of Lepidopsetta from other pleuronectid genera. Depending on developmental stage, larvae of Lepidopsetta generally are categorized by the presence of at least one postanal bar (Table 12). Larvae of L. bilineata have a postanal pigment pattern with four dorsal pigment patches (the posteriormost aligning with a ventral patch forming a caudal bar), larvae of L. mochigarei have three dorsal patches (the posteriormost aligns with a ventral pigment patch approximating a caudal bar), whereas larvae of L. polyxystra n. sp. have two dorsal pigment patches (the posteriormost aligns with a ventral pigment patch forming a bar located at about 2/3 of body length). Among other pleuronectid genera that have a postanal band and bar pattern, preflexion and early flexion larvae of Lepidopsetta may be distinguished by the presence of a series of postanal ventral midline melanophores and fewer pigment spots along the notochord tip (Table 12).

In general, larvae of L. bilineata closely resemble those of Psettichthys among other genera (Table 12). Both have alternating patches of pigment on dorsal and ventral body margins, although Psettichthys has a series of small patches on the dorsal and ventral finfold margins whereas L. bilineata has isolated patches along the finfolds. A series of postanal ventral melanophores is present in L. bilineata and absent in Psettichthys, in which postanal pigment is usually restricted to three or four large spots. Larvae of L. bilineata generally have much less pigment on the jaws and isthmus than do larvae of Psettichtys.

Size at stage of development varies among species (Table 13). Larvae of L. bilineata begin transformation at smaller lengths than larvae of L. polyxystra n. sp. or L. mochigarei. Larvae of L. bilineata also have a larger orbit, shorter snout-to-anus length, and a slenderer body (see Table 2).

The juvenile stages of most eastern North Pacific pleuronectids are poorly known. Traditional early life history descriptions typically describe developmental stages on the basis of eggs and larvae captured in plankton nets and usually do not include transitional larvae that undergo changes associated with a benthic existence. Plankton surveys routinely use nets that do not effectively sample bottom waters where many pleuronectid juveniles eventually settle.

Pleuronectid juveniles can be separated by several characters. Perhaps the most important character is the size at which transformation occurs, although data are scarce on when transformation is completed. Among taxa for which data are available, eastern North Pacific pleuronectids can be grouped into three categories based on approximate transformation sizes: [is less than] 15 mm SL, 15-30 mm SL, and [is greater than] 30 mm SL. Genera with at least one species transforming at sizes [is less than] 15 mm SL include Limanda, Platichthys, Pleuronectes, and Pleuronichthys (sensu Cooper and Chapleau, 1998). Genera with larvae that transform at much larger sizes ([is greater than] 30 mm SL) include Atheresthes, Embassichthys, Glyptocephalus, Hippoglossoides, Microstomus, and Reinhardtius. Larvae of Lepidopsetta generally undergo transformation between 15 and 25 mm SL. Other eastern North Pacific pleuronectids that undergo transformation at similar sizes include Acanthopsetta, Eopsetta, Hippoglossus, Isopsetta, Lyopsetta, Parophrys, and Psettichthys.

Characters that separate juveniles further include meristics of vertebrae, median fin elements, and gill rakers (although gill rakers may not be fully formed); mouth size; and the shape of the lateral line. Generally juveniles of Lepidopsetta have lower meristics than juveniles of Atheresethes, Eopsetta, Hippoglosssoides, and Hippoglossus. Juveniles of Lepidopsetta develop the highly arched lateral line during this stage (15-25 mm SL). A combination of mouth size, residual larval pigment on the blind side of early juveniles, lateral line shape, and, in larger juveniles, number of gill rakers may help to separate other similar-looking eastern North Pacific pleuronectids (Acanthopsetta, Isopsetta, Lyopsetta, Parophrys, and Psettichthys).

Identification of early life history stages of Lepidopsetta has been confused in the literature for many years. Hickman (1959) described egg and larval development of the sand sole, Psettichthys melanostictus, on the basis of material collected from Puget Sound, Washington. He illustrated six larvae identified as Psettichthys from specimens reared from eggs (his Figs. 1 and 2) and from larvae collected by net (his Figs. 3-6). His Figures 3 and 4 can be identified as L. bilineata from the alignment of dorsal and ventral midline melanophores on the postanal body.

In her extensive study of reproduction and development in North Pacific flatfishes, Pertseva-Ostroumova (1961) described the early life history stages of L. bilineata and L. mochigarei. The description of L. bilineata is based on material from collections off the Kuril Islands, off the eastern and western coasts of Kamchatka, and in the western Bering Sea. Larvae from these geographic areas are L. polyxystra n. sp.

Blackburn (1973) provided figures of two specimens of L. bilineata (his appendix Fig. C-2, A and B) in his survey of ichthyoplankton from Skagit Bay, Puget Sound. His discussion, however, is clearly based on the larvae of both eastern North Pacific species.

Garrison and Miller (1982) reviewed the reproductive characteristics of L. bilineata, but their sources either represent both species (Smith, 1936; Forrester, 1969; Forrester and Thompson, 1969) or L. polyxystra n. sp. alone (Shubnikov and Lisovenko, 1964; Shvetsov, 1979). Ahlstrom et al. (1984) correctly illustrated Psettichthys larvae but presented data and a figure for L. bilineata (their Fig. 351b) based on Figure 22 (4) of Pertseva-Ostroumova (1961), now identified as L. polyxystra n. sp. Okiyama (1988) also presented material for L. bilineata, based on Figure 23(1) of Pertseva-Ostroumova (1961), which therefore represents L. polyxystra n. sp.

Matarese et al. (1989) separated larvae of Lepidopsetta from Psettichthys. Their description and figures of L. bilineata are based on what we now refer to as L. polyxystra n. sp., whereas the descriptions and figures of “Lepidopsetta 2” are now referred to as L. bilineata. Charter and Moser (1996) presented figures of both Psettichthys melanostictus and L. bilineata but, because Pertseva-Ostroumova (1961) was cited as a source for L. bilineata, some descriptive data are suspect. Data for L. bilineata, particularly in the introductory tables, may be based in part on L. polyxystra n. sp. (see tables “Pleuronectidae,” pages 1370-1373, Charter and Moser, 1996).

Key to juveniles and adults [is greater than] 30 mm of species of Lepidopsetta

1a Preopercular pores 8-13; lateral-line pores 95-119; sum of scales above and below lateral line 91-103; supraorbital pores 1-3; total gill rakers on first arches [is less than or equal to] 10, on upper part of first arch [is less than or equal to] 3….. L. mochigarei

southern Sea of Okhotsk to Korea

1b Preopercular pores 5-7; lateral-line pores 70-91; sum of scales above and below lateral line 65-88; supraorbital pores 1-8; total gill rakers on first arch 6-14 … 2

2a Total gill rakers on first arch typically [is greater than or equal to] 10, on upper part of first arch [is greater than or equal to] 3; supraorbital pores 1-2, rarely 3-7; blind side creamy white……… L. polyxystra n. sp.

Puget Sound to Sea of Okhotsk

2b Total gill rakers on first arch typically [is less than or equal to] 10, on upper part of first arch [is less than or equal to] 3; supraorbital pores 3-8; blind side with extensive bright white highlights … L. bilineata

Baja California to southeastern Bering Sea

Lepidopsetto bilineata (Ayres, 1855a) Southern rock sole

Figs. 1-6, 10, 12, 14-18; Tables 2-11, 13-14

Platessa bilineata Ayres, 1855a:2 (original description, one specimen, apparently lost, sex and size unknown, fish markets of San Francisco Bay, California).

Platichthys umbrosus Girard, 1856:136 (original description, one specimen, apparently lost, sex unknown, ca. 190 mm TL, Cape Flattery, Washington).

Pleuronectes perarcuatus Cope, 1873:32 (original description, one specimen; holotype, ANSP 8725, sex undetermined, 108 mm SL, Gulf of Alaska, “Sitka” or “Unalaska”).

Lepidopsetta bilineata umbrosa (in part) Jordan and Evermann, 1898:2643 (new combination, “Puget Sound and northward”).

Lepidopsetta bilineata bilineata Taranets, 1937:144 (new combination, keys).

Pleuronectes bilineata Sakamoto, 1984a:99 (new combination, phylogenetics).

Neotype

CAS 42650, 1(207.6 mm), Calif., Gulf of the Farallones, Sta. F43(2)N, M. Moriguchi, August-September 1978.

Other material examined

A total of 380 adult and juvenile specimens, 22.7-426 mm, including the neotype listed above, was examined. Sixty-four larvae were examined.

Adults Bering Sea: UW 041696, 3(222-245 mm), N of Unimak I., 55.0213 [degrees] N, 164.6009 [degrees] W. Aleutian Islands: SIO 94-164, 1(167 mm) of 10(103-193 mm), Unimak Pass, 54 [degrees] 15.1’N, 165 [degrees] 57.6’W, 4 June 1994; UW 041695, 3(170-210), Aleutian Is., NE of Umnak I., 53.44 [degrees] N, 168.4944 [degrees] W; Gulf of Alaska: ABL uncat, 1(153 mm), Ursus Channel, San Fernando I., 1 November 1956; ABL 64-696, 1(138 mm), Gore Point, S of Kenai Peninsula, 28 July 1963; ABL 67-211, 4(197-234 mm), Auke Bay, 19.2 km NW of Juneau, 22 May 1962; ABL 71-5, 2(200.5-205.5 mm), Baranof I., Point Conclusion, 17 May 1971; OS 6447, 3(166.1-184.6 mm), 57 [degrees] 10’N, 151 [degrees] 40.5’W, 77 m, 20 September 1978; OS 2156, 1(75.0 mm), Revillagigedo I., Ward Cove, 26 July 1949; OS 2175, 1(102.4 mm), Revillagigedo I., Ward Cove, 26 August 1949; SIO 72-219, 1(251 mm), Lituya Bay, 9.3 km SW of Harbor Pt., 25 June 1961; SIO 72-258, 1(289 mm), Baranof I., Katlian Bay, 57 [degrees] 10’N, 135 [degrees] 20’W; SIO 72-225, 1(288 mm), 2 [degrees] E of Kodiak I., 6 May 1962; SIO 76-300, 1(187.5 mm) of 28(145-280 mm), Kodiak shelf, 57 [degrees] 40’N, 150 [degrees] 37’W, 23 June 1976; SIO 69-478, 3(112-170 mm) of 7(51-195 mm), Afognak I., Kitoi Bay, NE of Kodiak I., 16-21 April 1968; UW 040267, 1(330 mm), off Kodiak; UW 044021, 1(255 mm), 55.8088 [degrees] N, 158.7503 [degrees] W, 67 m, 10 June 1996; USNM 054286, 1(358 mm), Ketchikan, Sta. TT2120, RV Albatross; USNM 130739, 1(252 mm), Prince William Sound, Macleod Harbor, 16 March 1941; UW 01666, 1(132 mm), Alexander Archipelago, off Wrangell I., 1 December 1931; UW 008376, 1(240 mm), Ketchikan, 23 September 1949; UW 040902, 1(182 mm), Prince William Sound, 1989; UW 008292, 1(162 mm), Southeast Alaska, 26-31 March 1950; UW 018837, 2(222.5-230.5 mm), 11.7 km NNW of Triangle I., 50 [degrees] 58.1’N, 129 [degrees] 4.9’W, 19 August 1960; UW 40264, 1(342 mm), 54.039 [degrees] N, 165.8153 [degrees] W; UW 27679, 23(20.5-57.4 mm), Prince William Sound; UW 044010, 1(240 mm), 59.4918 [degrees] N, 151.6185 [degrees] W, 41 m, 27 February 1996; UW 044028, 1(292 mm), 59.4885 [degrees] N, 151.6062 [degrees] W, 30 m, 27 February 1996; UW 044025, 2(210-285 mm), Kachemak Bay, 12 May 1996; UW 040268, 1(290 mm), Kodiak I.; UBC 65-525, 1(290 mm), off Baranof I.; UW 044013, 7(287-426 mm), 56.5 [degrees] N, 153.5 [degrees] W, R. Bonaduhr; UW 041801, 6(270-335 mm), 54 [degrees] N, 160.74 [degrees] W, British Columbia: UW 083483, 33.5 mm, off Vancouver Island, 48 [degrees] 39.5’N, 125 [degrees] 55.2’W, 19 September 1977. SIO 63-202, 1(247 mm), Strait of Georgia, Fraser R.; USNM 31993, 1(158.5 mm), Carter Bay, June 1882; UBC 53-85, 1(172 mm), Vancouver I.; UBC 53-50, 1(57.3 mm), Vancouver I.; UBC 53-68, 1(105 mm), Vancouver I.; UBC 53-50, 1(57.3 mm), Vancouver I., Departure Bay, 49.2167 [degrees] N, 123.95 [degrees] W; UBC 53-301, 7(101.5-177 mm), Vancouver I., Baynes Sound; UBC 53-245, 1(144.8 mm), Vancouver I., Burrard Inlet; UBC 55-281, 1(32.9 mm), Vancouver I., Departure Bay, 49.2167 [degrees] N, 123.95 [degrees] W; UBC 56-8, 1(220.5 mm); UBC 55-496, 3(217-235 mm), Point Gray, 3.2 km SE of North Arm of Fraser R.; UBC 56-519, 5(205-233 mm), Vancouver fish docks; UBC 61-393, 3(59-88 mm), Bute Inlet; UBC 61-484, 2(145-167 mm), Vancouver I., Burrard Inlet, 25 October 1961; UBC 61-609, 9(38.5-123 mm), Bute Inlet; UBC 61-232, 2(245 mm and one head only), Hecate Strait; UBC 60-416, 1(205 mm), Queen Charlotte Is., Gillat Arm; UBC 63-910, 1(148 mm), Howe Sound; UBC 63-732, 10(68-140 mm), off Keats I.; UW 044026, 1(306.5 mm), 48.57723 [degrees] N, 124.7803 [degrees] W, 25 August 1995; UW 044027, 2(293-308 mm), 48.74507 [degrees] N, 125.9744 [degrees] W, 28 August 1995; UBC 65-676, 6(44-142 mm), Graham I., McIntyre Bay; UBC 62-93, 6(80-142 mm), Nass Bay; UBC 61-686, 1(62.8 mm), S of Vargas I., 5 July 1934; UBC 61-621, 1(50.5 mm), Snake I., near Nanaimo, 8 May 1933; UBC 61-610, 2(139-142 mm), Work Channel, head of Trail Bay, 16 July 1951; UBC 61-674, 1(60 mm), Dean Channel, off Nescall Bay; UBC 62-94, 1(94 mm), Sydney Inlet; UBC 81-3, 1(201 mm), Vancouver I., Grappler Inlet; UBC 76-7, 1(135 mm), English Bay, 20 January 1962. Washington, Oregon, and California: USNM 054392,1(292 mm), near Port Townsend, TT2838, RV Albatross; USNM 27299, 2(150-246 mm), Puget Sound, 1880; UW 041090, 2(196.0-196.5 mm), Agate Pass, 3 April 1950; UW 041089, 1(229 mm), Agate Pass, 3 April 1950; UW 6070, 7(61-133 mm), San Juan Islands, East Sound, 3 March 1937; UW 06134, 2(53-63.5 mm), N of Maury I., 4 December 1948; UW 014838, 2(65-68 mm), Golden Gardens Beach, 14-15 May 1952; UW 014380, 2(101-108 mm), near Hat I., 29 May 1936; UW 044011, 1(145 mm), San Juan Islands, East Sound, 18 February 1964; UW 017115, 1(1178 mm), Port Orchard Channel, 2 February 1964; UW 015423, 1(169 mm), Shilshole Bay; UW 000782, 2(147-156.5 mm), Alki Point, 8 March 1930; UW 005522, 11(46.6-159.5 mm), Edmonds, 18 August 1947; UW 018663, 1(60.5 mm), San Juan Islands, East Sound, 11 December 1964; UW 040665, 2(54-58 mm), Port Townsend Bay, 5 January 1979; UW 040686, 6(114.3-167.3 mm), West Point, 3 October 1978; UW 025723, 4(52.1-230.5 mm), Port Townsend Bay, 5 December 1978-5 January 1979; UW 025721, 3(166-208 mm), Golden Gardens, 22 July 1981; UW 006109, 1(154.5 mm), Alki Point, 3 March 1939; UW 017840, 1(211 mm), S end of Port Susan, 6 May 1950; UW 025719, 12 (103.2-164.5 mm), Murden Cove, 22 April 1980; UW 025727, 2(116.5-169.5 mm), West Point, 29 March 1979; UW 005135, 8(22.7-70.6 mm), Alki Point, 5 April 1938; UW 025197, 1(77.3 mm; cleared-and-stained), Meadow Point; UW 044006, 11(135-250.5 mm), Nisqually, 47.1578 [degrees] N, 122.6693 [degrees] W, 16 May 1996; UW 040269, 10(209-259 mm), Puget Sound, 47 [degrees] 19.48’N, 122 [degrees] 33.8’W, 10 May 1996; UW 029670, 49(125.3-283.5 mm), Colvos Passage off Vashon I., 47.5 [degrees] N, 122.4167 [degrees] W, 18 July 1949; UW 025726, 5(125-175 mm), Puget Sound, West Point, 3 October 1978; UW 025720, 7(130-185 mm), Puget Sound, West Point, 3 April 1979; UW 041319, 6(122-202 mm), Puget Sound, West Point; UW 041330, 1(225 mm), Puget Sound, Portage Bay; UW 041301, 1(245 mm), Puget Sound, Port Townsend Bay; OS 15509, 4(226.5-287.4 mm), Cobb Seamount, 46 [degrees] 50’N, 130 [degrees] 50’W, 20 August 1992; OS 13792, 1(180.3 mm), 44 [degrees] 54.2667’N, 124 [degrees] 10.0667’W; OS 13500, 1(417 mm), Cobb Seamount, 46 [degrees] 50’N, 130 [degrees] 50’W; OS 13499, 1(330 mm), 44 [degrees] 52.2667’N, 124 [degrees] 09.55’W; OS 7482,1(223.3 mm), off Newport; OS 14732, 1(132.2 mm), 44 [degrees] 40.7’N, 124 [degrees] 09’W; OS 14690, 1(189.4 mm), 44 [degrees] 37’N, 124 [degrees] 11.1’W; OS 6208, 4(137.3-222.8 mm), off Newport, 30 September 1978; OS 7477, 1(173.7 mm), 44 [degrees] 37.4’N, 124 [degrees] 08.9’W; CAS 18553, 1(198.5 mm), Calif., San Francisco Bay, 14 May 1931; CAS 31832, 1(205 mm), Ca]if., Gulf of the Farallones, 37 [degrees] 44’N, 122 [degrees] 40’W, October 1973; CAS 40341, 1(347.4 mm), Calif., 22.6 km NW of Pt. Pinos, 34 [degrees] 47.5’N, 122 [degrees] 7.8167’W; CAS-SU 111615, 1(335.0 mm), San Francisco Market, May 1895; CAS 49152, 1(229.5 mm), Calif., Sonoma, Bodega Bay, 18 October 1981; SIO 88-119, 1(277 mm), Calif., south of La Jolla; SIO H48-22, 1(261 mm), Calif., south of La Jolla; SIO 63-241, 1(243 mm), Tanner Bank, Calif., 32 [degrees] 42.5’N, 119 [degrees] 6.5’W, 17 May 1963; SIO 65-460, 1(284 mm), Calif., ridge between Tanner and Cortez Banks, 26 September 1965; SIO 65-6-64A, 1(210 mm), Calif., Tanner Bank, 32 [degrees] 42’N, 119 [degrees] 08’W, 15 January 1965; SIO 63-732-64A, 1(183 mm), Calif., Catalina I., off Avalon, 22 March 1962; SIO 85-145, 2(74-86 mm), Calif., Navarro Head, 39 [degrees] 05.7’N, 123 [degrees] E44.95’W, 60.4-110.5 m, 17 August 1985; LACM W54-382, 7(178.5-247 mm), 1.6 km W of Santa Barbara I., 33 [degrees] 08.15’N, 119 [degrees] 04.1333’W, 31 October 1954; LACM W66-67, 1(198.5 mm), off San Simeon Point, 13 November 1966; LACM W54-380, 2(274-275 mm), 1.6 km E of Santa Barbara I., 30 October 1954; LACM 44390-1, 1(282 mm), off Palos Verdes, 22 February 1988; LACM 31966-8, 8(176.5-357 mm), Humboldt Co., 8 km S of Shelter Cove, 7 August 1971; LACM 241, 4(153-320 mm), Calif., San Francisco; OS 972,1(201.2 mm), southern Calif.; LACM 35690-3, 2(83-164.5 mm), Calif., S end of Tanner Bank; UW 044009, 9(143-252.5 mm), 37.781 [degrees] N, 122.9288 [degrees] W, 26 June 1995; UW 040262, 1(250 mm), 48.74317 [degrees] N, 125.7785 [degrees] W, 27 September 1992; UW 041698, 1(223 mm), 35.0935 [degrees] N, 120.7774 [degrees] W, 15 June 1995; UW 041803, 1(358 mm), 47.0839 [degrees] N, 124.748 [degrees] W, 8 August 1995; UW 041802, 1(376 mm), 46.91147 [degrees] N, 124.7292 [degrees] W, 3 August 1995; UW 041903, 2(290-347 mm), 46.91308 [degrees] N, 124.4756 [degrees] W, 5 August 1995; Mexico: CAS 2484, 1(226.3 mm), Baja California, off Cortez Banks, 21 August 1932.

Larvae 64 specimens (2.7-19.0 mm) examined: Western Gulf of Alaska: UW 083400, 2(10.8-12.3 mm), 54 [degrees] 10.8’N, 165 [degrees] 24.3’W, 0-55 m depth, Methot net, 30 July 1991; UW 083401, 1(10.9 mm), 54 [degrees] 38.5’N, 160 [degrees] 51.2’W, 0-98 in depth, Methot net, 30 July 1991; UW 083402, 1(12.0 mm), 56 [degrees] 11.6’N, 157 [degrees] 21.7’W, 0-129 m depth, Methot net, 27 July 1991; UW 083403, 1(14.0 mm), 54 [degrees] 36.2’N, 162 [degrees] 16.2’W, 0-87 m depth, Methot net, 30 July 1991; UW 083404, 1(14.1 mm), 55 [degrees] 43.8’N, 157 [degrees] 34.6’W, 0-105 m depth, Methot net, 27 July 1991; UW 083405, 2(14.4-15.6 mm), 55 [degrees] 43.0’N, 159 [degrees] 49.4’W, 0-108 m depth, Methot net, 25 July 1991; UW 083406, 1(15.5 mm), 53 [degrees] 59.8’N, 165 [degrees] 51.0’W, 0-50 m depth, Methot net, 31 July 1991; UW 083407, 2(15.9-16.0 mm), 55 [degrees] 18.0’N, 160 [degrees] 11.4’W, 0-115 m depth, Methot net, 24 July 1991; UW 083408,1(16.0 mm), 55 [degrees] 17.7’N, 160 [degrees] 11.7’W, 0-115 m depth, Methot net, 24 July 1991; UW 083445 1(13.3 mm), 56 [degrees] 11.6’N, 158 [degrees] 03.6’W, 0-141 m depth, Methot net, 25 July 1991; UW 072255, 5 of 6(2.7-8.6 mm), 57 [degrees] 01.1’N, 156 [degrees] 19.2’W, 0-95 m depth, bongo net, 3 June 1990; UW 072453, 2 of 4(6.2-6.5 mm), 57 [degrees] 17.2’N, 155 [degrees] 27.6’W, 0-102 m depth, bongo net, 5 June 1990; UW 072126, 2(6.6-7.3 mm), 56 [degrees] 54.0’N, 156 [degrees] 29.5’W, 0-84 m depth, bongo net, 3 June 1990; UW 071455, 1(6.7 mm), 57 [degrees] 60.0’N, 153 [degrees] 59.3’W, 0-103 m depth, bongo net, 28 May 1990; UW 072087, 1(8.1 mm), 56 [degrees] 39.8’N, 156 [degrees] 20.0’W, 0-102 m depth, bongo net, 3 June 1990; UW 072264, 1(8.7 mm), 57 [degrees] 09.1’N, 156 [degrees] 09.1’W, 0-100 m depth, bongo net, 3 June 1990; Gulf of Alaska: UW 083484, 1(19.0 mm), 59 [degrees] 37.8’N, 151 [degrees] 44.2’W, 34 m depth, 17 July 1996; UW 083409, 1(16.4 mm), 58 [degrees] 19.5’N, 150 [degrees] 53.0’W, 64 m depth, bongo net, 22 July 1977; UW 083410, 1(6.3 mm), 57 [degrees] 61.1’N, 151 [degrees] 17.4’W, 0-35 m depth, Tucker net, 26 June 1978; UW 083411, 1(9.7 mm), 56 [degrees] 42.3’N, 153 [degrees] 33.4’W, 28-70 m depth, Tucker net, 28 June 1978; UW 083412, 1(11.0 mm), 57 [degrees] 00.8’N, 153 [degrees] 28.3’W, 0 m depth, neuston net, 14 September 1978; UW 083413, 2(5.1-6.2 mm), 57 [degrees] 10.7’N, 156 [degrees] 01.3’W, 0-100 m depth, Tucker net, 4 June 1988; UW 083414, 1 (6.6 mm), 56 [degrees] 21.0’N, 157 [degrees] 03.8’W, 0-100 m depth, Tucker net, 6 June 1988; UW 083415, 1(6.4 mm), 56 [degrees] 47.3’N, 155 [degrees] 26.0’W, 0-105 m depth, Tucker net, 23 May 1988; UW 083416, 1(6.5 mm), 57 [degrees] 03.8’N, 156 [degrees] 01.8’W, 0-101 m depth, Tucker net, 4 June 1988; UW 083417, 1(6.6 mm), 56 [degrees] 47.3’N, 155 [degrees] 25.9’W, 0-101 m depth, Tucker net, 4 June 1988; UW 083418, 1(7.1 mm), 57 [degrees] 10.8’N, 156 [degrees] 29.9’W, 0-102 m depth, Tucker net, 4 June 1988; UW 083419, 1(8.3 mm), 56 [degrees] 38.9’N, 156 [degrees] 31.5’W, 0-101 m depth, Tucker net, 2 June 1988; UW 083420, 1(10.2 mm), 56 [degrees] 46.7’N, 156 [degrees] 18.3’W, 0-91 m depth, Tucker net, 2 June 1988; UW 083421, 1(10.6 mm), 57 [degrees] 15.0’N, 155 [degrees] 53.6’W, 0-101 m depth, Tucker net, 4 June 1988; UW 083422, 3(3.3-3.4 mm), 57 [degrees] 55.6’N, 151 [degrees] 02.3’W, 0-76 m depth, bongo net, 13 May 1991; UW 083423, 1(3.9 mm), 5 [degrees] 43.8’N, 154 [degrees] 02.1’W, 0-100 m depth, bongo net, 24 May 1991; UW 083424, 1(4.7 mm), 55 [degrees] 53.9’N, 155 [degrees] 59.8’W, 0-75 m depth, bongo net, 23 May 1991; UW 083425, 1(4.9 mm), 57 [degrees] 36.9’N, 155 [degrees] 28.3’W, 0-101 m depth, bongo net, 24 May 1991; UW 083426, 1(3.6 mm), 56 [degrees] 40.TN, 155 [degrees] 10.7’W, 0-61 m depth, bongo net, 8 May 1992; UW 083427, 1(6.0 mm), 56 [degrees] 58.4’N, 156 [degrees] 06.2’W, 0-100 m depth, bongo net, 14 May 1992; UW 083428, 2(3.5-4.3 mm), 55 [degrees] 27.2’N, 157 [degrees] 39.3’W, 0-88 m depth, bongo net, 19 May 1992; UW 083429, 1(3.8 mm), 55 [degrees] 17.0’N, 157 [degrees] 44.8’W, 0-72 m depth, bongo net, 19 May 1992; UW 083430, 1(4.6 mm), 55 [degrees] 22.5’N, 156 [degrees] 56.8’W, 0-80 m depth, bongo net, 19 May 1992; UW 083431, 1(7.9 mm), 55 [degrees] 16.0’N, 156 [degrees] 15.0’W, 0-102 m depth, bongo net, 18 May 1992; UW 083432, 1(7.3 mm), 55 [degrees] 55.3’N, 156 [degrees] 15.3’W, 0-216 m depth, bongo net, 21 May 1992; UW 083443, 1(3.2 mm), 57 [degrees] 01.1’N, 156 [degrees] 19.2’N, 0-95 m depth, bongo net, 3 June 1990; Puget Sound: UW 083433, 1(9.6 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.3’W, 0 m depth, dip net, 9 May 1994; UW 083434, 1(9.7 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.3’W, 0 m depth, dip net, 5 May 1989; UW 083435, 1(11.2 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.3’W, 0 m depth, dip net, 27 March 1991; UW 083436,1(11.4 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.3’W, 0 m depth, dip net, 16 July 1987; UW 083437, 1(13,3 mm), 48 [degrees] 1.0’N, 123 [degrees] 0.0’W, reared, 4 May 1989; UW 083438, 1(13.4 mm), 48 [degrees] 1.0’N, 123 [degrees] 0.0’W, reared, 4 May 1989; UW 083439, 1(14.4 mm), 48 [degrees] 1.0’N, 123 [degrees] 0.0’W, 0 m depth, dip net (26 April 1989), reared (20 May 1989); UW 083440, 1(15.2 mm), 48 [degrees] 1.0’N, 123 [degrees] 0.0’W, 0 m depth, dip net (26 April 1989), reared (20 May 1989); UW 083441, 1(15.6 mm), 48 [degrees] 1.0’N, 123 [degrees] 0.0’W, 0 m depth, dip net, 1 July 1989; UW 083442, 1(11.4 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.3’W, 0 m depth, dip net, 16 July 1987; UW 083444, 1(9.3 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.3’W, 0 m depth, dip net, 1 June 1989.

Diagnosis

This species of Lepidopsetta has the following combination of characters in adults: total gill rakers on first arch 6-11; on upper arch 1-3, rarely 4, with at least one rudimentary; total gill rakers on second arch 6-11; supraorbital pores 3-9; preopercular pores 5-7; lateral-line pores 70-91; sum of scales above and below lateral line 65-88; interorbital narrow; blind-side coloration white, with glossy highlights along myotome margins increasing anteriorly.

Larvae are distinguished from other species of Lepidopsetta by the following characters: body deep, snout-to-anus length short; hatching, flexion, and transformation at comparatively smaller sizes; preflexion pigment pattern with pigment patches along distal edges of dorsal and anal finfolds, four prominent spots along dorsal midline (in a pattern resembling alternating dorsal and ventral spots, with posteriormost dorsal spot coalescing with a corresponding ventral patch to form a bar), and a series of small ventral midline melanophores extending from gut to last myomere; flexion pigment pattern with distinctive anterior dorsal midline spot and posteriormost dorsal spot forming a bar with corresponding ventral patch; pectoral-fin rays unpigmented.

Description of adults (Fig. 15)

[Figure 15 ILLUSTRATION OMITTED]

Body ovate, greatest depth 40.8-57.0 (47.0)% SL, scales above lateral line 25-43 and scales below lateral line 27-53; head relatively acute, length 24.0-32.9 (28.2)% SL; dorsal margin of head at dorsal-fin origin concave, snout length 3.1-6.1 (4.5)% SL (12.4-22.4 (16.1)% HL); ocular-side maxilla length 25.3-32.5 (27.9)% HL; blindside maxilla relatively long, length 25.2-41.2 (31.3)% HL; ocular-side mandible length 38.3-46.1 (41.9)% HL; teeth 5-10 on ocular-side premaxilla, 20-27 on blind-side premaxilla and 10-12 on ocular-side dentary, 23-29 on blindside dentary; gill rakers of first arch typically broad and robust, 6-11 total, 1-4 on upper arch, 5-7 on lower; gill rakers of second arch 6-11 total, 1 on upper and 5-10 on lower arch; dorsal orbit larger than eye length, orbit length 23.4-32.5 (28.1)% HL, dorsal eye length 17.9-30.3 (24.2)% HL; ventral eye length 18.2-35.4 (24.8)% HL; interorbital narrow, up to 3 scales at narrowest portion, 2.1-6.0 (3.5)% HL; cheek with 9-16 scales, length 28.1-41.8 (34.2)% HL, depth 13.9-24.2 (19.2)% HL; preopercular pores 5-7; ocular-side suborbital pores 14-29; blind-side suborbital pores 6-13; lateral-line pores 70-91, lateralline arch length 49.3-58.1 (54.3)% HL, its depth 24.8-37.1 (30.2)% of its length; both anterior and posterior supratemporal branches relatively long, anterior pores 2-12, posterior pores 8-30; supraorbital canal long, extending to dorsal rim of dorsal orbit near insertions of dorsal-fin rays 3-4, pores 3-9; ocular-side pectoral-fin length 12.2-18.2 (15.2)% SL (43.6-66.2 (53.9)% HL); blind-side pectoral-fin length 6.7-12.2 (9.9)% SL (23.2-44.3 (35.0)% HL), about equal to ocular-side pelvic-fin length 7.6-12.5 (9.8)% SL (26.7-44.7 (34.8)% HL); dorsal fin with 67-89 rays, height 10.7-16.5 (13.3)% SL, in specimens with 72-79 rays supported by 71-78 pterygiophores, 9-11 anterior to first neural spine; anal fin with 54-77 rays, in specimens with 54-62 rays supported by 53-60 pterygiophores; caudal peduncle relatively slender, least depth 8.2-12.3 (10.2)% SL (29.6-43.0 (36.2)% HL; 90.0-152.4 (112.6)% caudal peduncle length); greatest depth 10.5-14.3 (12.1)% SL (37.2-53.0 (42.8)% HL); caudal peduncle length 7.2-10.9 (9.1)% SL (25.5-42.2 (32.4)% HL); caudal-fin length 15.3-23.6 (20.8)% SL. Vertebrae 40-41, with 11 precaudal and 29-30 caudal.

Scales around head and those scattered posteriorly on ocular side moderately rough with columnar tubercles in large adults; strong spines in small adults and juveniles. Urogenital flap darkly pigmented in 21-58 mm juveniles.

In life, blind side in adults translucent to bright white, with glossy highlights along edges of myotomes; in juveniles, primarily translucent, and having reduced glossy areas especially prominent over the head. When preserved, blind side of all individuals uniform creamy white to yellow-brown. Ocular side slightly more green than that of congeneric in Puget Sound.

Remaining description as for genus. Largest specimen examined 426 mm (UW 044012). Maximum size reported ca. 540 mm (580 mm TL(6)).

Description of juveniles (Fig. 16)

[Figure 16 ILLUSTRATION OMITTED]

Most individuals were collected on bottom (Fig. 16A) by 19.0 mm; newly settled juveniles with dorsal eye completely migrated; median fin rays formed; lateral line nearly formed; expanded anteriormost anal-fin pterygiophore well developed. Preanal pigment increased along head and gut; postanal pigment smaller and distributed in random patches; urogenital papilla darkly pigmented throughout its length. Blind side with pigment similar to that of postflexion larvae. Orbit length larger, mouth smaller, body depth less, gill-raker counts on lower arch less, and distance from pelvic-fin origin to anal-fin origin greater than in L. polyxystra n. sp. Pelvic- and pectoral-fin rays formed.

By 30.0 mm, pigmentation has increased along body, with darker patches and spots throughout (Fig. 16B) obscuring pigmented urogenital papillae. Lateral line more fully developed; supraorbital canal pores visible.

Description of larvae (Fig. 17)

[Figure 17 ILLUSTRATION OMITTED]

Snout-to-anus length is 32.934.6% SL, remaining constant during development; body depth 4.7-35.7% SL, increasing with development, sharply after flexion; head length 13.3-29.3% SL, increasing with development, sharply after flexion; snout length 22.6-20.7% HL, remaining constant during development; orbit length 51.9-23.8% HL, decreasing with development (Table 2). Total myomeres 37-44.

Larvae hatching at small lengths, at sizes less than 3.0 mm, yolk absorbed by 2.7-4.5 mm. Preflexion larvae ranging in size from 3.8 to 6.6 mm; flexion larvae, from 6.4 to 11.1 mm; postflexion larvae from 10.8 to 16.0 mm. Transformation occurring at lengths as small as 10.0 mm (often accompanied by a decrease in total body length); postsettlement juvenile stage usually attained by 20.0 mm (Table 13).

Preanal pigment present initially along lower jaw and ventral side of cleithral region, increasing with development to snout, upper jaw, and isthmus. Pigment ventrally along gut and dorsally on anus; by flexion a distinct patch of melanophores along the posterior edge of the gut; pigment increases laterally with development.

Postanal pigment present as melanophores along distal edges of dorsal and anal finfolds; four distinct pigment areas along the dorsal midline, anterior (first) spot begins 1-5 myomeres after anus at about myomere 12-16, second spot begins at about myomere 23-26, third spot begins at about myomere 33-46, and the fourth spot begins at about myomere 41-42 (after initially forming as a dorsal midline patch, the posteriormost spot coalesces with ventral patch to form a caudal bar); series of melanophores from the gut along the ventral midline beginning in a double row, changing to a single row posterior to the ventral stripe of the caudal bar; several additional melanophores along the ventral midline posterior to the caudal bar, pigment above and below the tip of the notochord. By transformation, the third dorsal midline patch and opposing ventral patch expand to form an indistinct bar; other patches of pigment form in myosepta and continue into the dorsal and anal pterygiophores and fin rays.

Distribution (Figs. 6, 10, 12, 18)

Lepidopsetta bilineata ranges from the continental shelf north and south of the Islands of Four Mountains in the eastern Aleutian Islands and in the southern Bering Sea on the Slime Bank north of Unimak I., to Cortez Banks, Baja California, Mexico. It is common from the northern Gulf of Alaska to Puget Sound and is locally abundant along the coasts of Washington, Oregon, and California. Larvae have been collected from just south of the Aleutian Islands to Tanner Bank, Mexico (Moser et al., 1993; Charter and Moser, 1996).

Habitat

Adults were collected over sand and gravel substrates to depths of 339 m (RACE(6)). Larvae were collected over depths [is less than] 1000 m.

During our 22-year sampling period, larvae were less common than those of L. polyxystra n. sp. in spring ichthyoplankton surveys conducted in the Gulf of Alaska. Larvae appeared in larger numbers later in the season (June) and the highest densities occurred from Kodiak Island to the eastern Gulf of Alaska (Table 14; Fig. 18). In the CalCOFI region, larvae were collected from February to July; peak abundance was in May (Charter and Moser, 1996). Larvae were collected more frequently and in higher densities closer to the coastline (within 55.6 km, Moser et al., 1993).

[Figure 18 ILLUSTRATION OMITTED] Life history

In L. bilineata taken off the coast of Oregon, brittlestars of the genus Ophiura dominated the diet, and polychaetes and mollusks constituted much of the remainder (Kravitz et al. 1976). Adults from the Gulf of Alaska and Aleutian Islands are often infested with the parasitic copepods Nectobrachia indivisa and Naobranchia occidentalis.(7) Both parasites were previously recorded in Lepidopsetta by Kabata (1988). For both parasite species, L. bilineata was significantly less infested than L. polyxystra n. sp.(6) The maximum recorded age for female Lepidopsetta is 18 yr at 49 cm FL and for a male is 17 yr at 40 cm FL (Levings, 1967; Forrester, 1969). For early life history information, see generic account.

Etymology

The specific name is derived from the Latin bilineata, meaning “two lined”, a reference to the bifurcate supratemporal branch of the lateral line.

Comments

Basing his work on an account first published in The Pacific (Ayres, 1855a) and subsequently reprinted in the Proceedings of the California Academy of Sciences (Ayres, 1855b), Ayres described his specimen of Platessa bilineata at the California Academy of Natural Sciences (CAS) in 1855. The specimen has not been located at CAS(8) (where it may have been destroyed in the 1906 San Francisco earthquake and subsequent fire, Springer and Anderson, 1998), ANSP,(9) LACM,(10) or USNM.(11) Because L. bilineata may be easily confused with L. polyxystra n. sp., especially in the Gulf of Alaska and Puget Sound, we therefore designated CAS 42650, collected in the Gulf of Farallones off California, as the neotype of L. bilineata. Ayres obtained his specimen from the San Francisco fish market and noted that, although not common, the species was taken in San Francisco Bay. Lockington (1879a) noted that L. bilineata was most commonly caught “at or near the Farallone Islands” for the San Francisco markets.

Girard (1856) described Platichthys umbrosus on the basis of a single specimen deposited at the USNM, which was collected at Cape Flattery, Washington, by Lieutenant W. P. Trowbridge. One lot listed in the catalog, USNM 607, bears the collector’s name as Trowbridge and a collection date of 1856, and thus the single specimen of this lot appears to have been the one examined by Girard for his original description. From the original description, the specimen is approximately 190 mm TL. Unfortunately, this lot cannot be found and is presumed missing or destroyed.(11)

Girard was apparently unaware of Ayres’ (1855a, 1855b) description of L. bilineata (Leviton and Aldrich, 1997). Although Gunther (1862) considered Platichthys umbrosus distinct (basing his conclusion on the dried skins identified as P. umbrosus taken off Vancouver Island, Canada, and on the descriptions of L. bilineata provided by Ayres), later authors have treated P. umbrosus as a synonym of L. bilineata (Lockington, 1880a, 1880b; Jordan and Gilbert, 1881; Jordan and Evermann, 1898; Norman, 1934; Schmidt, 1950), and the name has not been used at the species level since Lockington’s (1879b) misidentification of Isopsetta isolepis. Lepidopsetta polyxystra n. sp. also ranges south along the east coast of Vancouver Island, the “Inside Passage,” to Puget Sound, and larvae of the new species have been recorded from the extreme northwest coast of Vancouver Island. Although L. bilineata is present, but uncommon, off the west coast of Vancouver Island, no adults or larvae of L. polyxystra n. sp. have been collected off Washington (Cape Flattery and south), despite the intensive survey efforts of the 1995 and 1998 NMFS triennial surveys, during which all captured Lepidopsetta were examined. Although the original description is insufficient to refer it to either of these species, the type locality lies well within the known range of L. bilineata and outside that of L. polyxystra n. sp. and we therefore consider Platichthys umbrosus to be a synonym of L. bilineata.

Cope’s (1873) description of Pleuronectes perarcuatus, from a specimen taken somewhere along the eastern Pacific Ocean coast of Alaska, perhaps near Sitka or Unalaska Island, included a comparison with Girard’s Platichthys umbrosus but not with Ayres’s Platessa bilineata. Although the holotype is badly damaged, especially about the head, the gill-raker count and structure, as well as presence of at least one supraorbital pore at the dorsal margin of the orbit,(8) indicate the holotype is L. bilineata. The specimen has a high lateral-line pore count at the upper range of counts for L. bilineata. The name has been considered synonymous with, or a subspecies of, L. bilineata by later authors (Jordan and Gilbert, 1881; Jordan and Goss, 1889; Jordan and Evermann, 1898; Wilimovsky et al., 1967).

Lepidopsetta mochigarei Snyder, 1911 Ricecake sole Asabagarei Figs. 1-2, 7, 19-20; Tables 3-10, 13

[Figures 19-20 ILLUSTRATION OMITTED]

Lepidopsetta mochigarei Snyder, 1911:547 (original description, two specimens: holotype by original designation, USNM 68245, male, 174 mm, and “cotype.” SU 21430, 145 mm TL, both from the market at Otaru, Hokkaido, Japan).

Lepidopsetta bilineata mochigarei Taranets, 1937:144 (new combination, keys).

Pleuronectes mochigarei Sakamoto, 1984a:99 (new combination, phylogenetics).

Material examined

Adults 27 specimens, 90.7-280.3 mm, including holotype listed above. Japan: CAS-SU 113378, 1(257.5 mm), Bomasiri Shima, off N end of Rebun To., 45 [degrees] 25.5’N, 140 [degrees] 53’E, 22 September 1906; CAS-SU 122548, 1(146.7 mm), Tsugaru Strait/Sea of Japan, between Hakodate, Hokkaido and Ebisu, Sado I., S of Cape Tsiuka, 41 [degrees] 35.8333’N, 140 [degrees] 36.75’E, 16 July 1906; HUMZ 56717, 1(154 mm), Hokkaido, off Kushiro; HUMZ 58576, 1(189 mm), Hokkaido, off Otanoshike; HUMZ 58566, 1(197.5 mm), Otanoshike, off Hokkaido, 20 m; HUMZ 58563, 1(200.5 mm), Otanoshike, off Hokkaido, 20 m; HUMZ 58568, 1(171.5 mm), Otanoshike, off Hokkaido; HUMZ 58567, 1(184 mm), Hokkaido, off Otanoshike; HUMZ 87820, 1(175 mm), Hokkaido, off Muroran, 135 m; HUMZ 80974, 1(190.5 mm), Hokkaido, off Muroran; HUMZ 81056, 1(167.5 mm), Hokkaido, Usujiri, Minamikayabe; HUMZ 90912, 1(142.5 mm), Hokkaido, off Tomakomai, 300 m; HUMZ 64730, 1(116.6 mm), Hokkaido, Funkawan; HUMZ 15505, 1(107.2 mm), Hokkaido, Funka Bay; HUMZ 15456, 1(130.5 mm), Hokkaido, Funka Bay; HUMZ 15506, 1(90.7 mm), Hokkaido, Funka Bay; HUMZ 94034, 1(193.5 mm), Hokkaido, off Tomakomai, 300 m; UBC 58-355, 2(204-240 mm), Japan; USNM 150380, 1(130.6 mm), Hokkaido; USNM 077125, 1(166.1 mm), 104 km S of OseSaki, Albatross Sta. 5067, 35.0972 [degrees] N, 138.6875 [degrees] E, 536 m; Russia: USNM 77126, 1(244.2 mm), Gulf of Tartary, off SW coast of Sakhalin I., Albatross Sta. 4999, 47.6389 [degrees] N, 141.65 [degrees] E; USNM 77127, 1(247.4 mm), Aniwa Bay, approaching Korsokov, Sakhalin I., 73 m, Albatross Sta. 5008, 46.1306 [degrees] N, 142.6222 [degrees] E; USNM 77130, 1(280.3 mm), Gulf of Tartary, off SW coast of Sakhalin I., 57 m, Albatross Sta. 5000, 46.55 [degrees] N, 142.7083 [degrees] E; USNM 77128, 1(257 mm), Aniwa Bay, approaching Korsokov, Sakhalin I., 77 m, Albatross Sta. 5007, 46.05 [degrees] N, 142.5167 [degrees] E; USNM 77135, 1(246.8 mm), Sakhalin I., off Korsokov Lt., 39-56 m, Albatross Sta. 5010, 46.55 [degrees] N, 142.725 [degrees] E.

Diagnosis

Lepidopsetta mochigarei is a species of Lepidopsetta with the following combination of characters in adults: total gill rakers on first arch 6-10, on upper arch 1-3; total gill rakers on second arch 7-10; supraorbital pores 1-3; preopercular pores 8-13; lateral-line pores 95-119; sum of scales above and below lateral line 91-108; interorbital narrow; blindside coloration white, and glossy highlights along myotome margins increasing anteriorly.

Larvae are distinguished from other species of Lepidopsetta by having the following combination of characters: body deep, increasing rapidly with development and snout-to-anus length moderate; larvae undergoing notochord flexion at comparatively larger sizes; preflexion pigment pattern with three prominent spots along margin of dorsal finfold and two spots along ventral finfold, three patches along postanal dorsal midline aligned with finfold spots to form slightly offset opposing patches, posteriormost dorsal spot coalescing with a corresponding ventral spot to form a bar; pectoral-fin rays pigmented (Okiyama and Takahashi, 1976; Nagasawa, cited in Okiyama, 1988).

Description of adults

Body ovate, relatively deep, greatest depth 45.5-61.1 (51.7)% SL, scales above lateral line 35-45 scales below lateral line 52-67; head relatively short and robust, length 23.3-30.5 (26.2)% SL; dorsal margin of head at dorsal-fin origin nearly linear, snout length 3.3-6.2 (4.3)% SL (12.7-20.2 (16.1)% HL); ocular-side maxilla length 24.6-29.4 (27.2)% HL; blind-side maxilla length 27.3-32.9 (29.7)% HL; ocular-side mandible length 36.6-44.6 (39.7)% HL; teeth 6-10 on ocular-side premaxilla, 23 on blindside premaxilla and 8-15 on ocular-side dentary, 21-28 on blind-side dentary; gill rakers of first arch broad and robust, 6-10 total, 1-3 on upper arch, 4-7 on lower; gill rakers of second arch 7-10 total, 1 on upper and 6-9 on lower arch; eyes relatively large, dorsal orbit slightly longer than eye length, orbit length 26.1-34.7 (30.1)% HL, eye length 21.4-30.0 (25.3)% HL; ventral eye length 21.6-31.1 (25.5)% HL; interorbital a narrow ridge, up to 2 scales at narrowest portion, 1.3-4.2 (2.7)% HL; cheek with 10-15 scales, length 28.6-35.0 (31.1)% HL, depth 14.8-25.5 (19.7)% HL; preopercular pores 8-13; ocularside suborbital pores 22-38; blind-side suborbital pores 10-20; lateral line pores 95-119, lateral-line arch relatively long, 53.7-69.9 (62.7)% HL, its depth 23.8-35.7 (30.9)% its length; both anterior and posterior supratemporal branches relatively short, anterior pores 3-12, posterior pores 6-22; supraorbital canal short, reaching only to the posterior rim of the dorsal orbit at about the insertion of dorsal-fin rays 5-6, pores 1-3; ocular-side pectoral-fin length 12.6-17.4 (14.6)% SL (47.9-60.2 (55.6)% HL); blind-side pectoral-fin length 7.9-11.7 (9.6)% SL (30.3-42.8 (36.5)% HL); ocular-side pelvic-fin length 7.4-11.8 (9.8)% SL (31.8-43.5 (37.4)% HL); dorsal fin with 73-82 rays, height 11.2-14.7 (12.9)% SL, in specimens with 75-81 rays supported by 71-77 pterygiophores, typically 8 or rarely 9 anterior to first neural spine; anal fin with 56-65 rays, in specimens with 56-63 rays supported by 57-61 pterygiophores; caudal peduncle relatively deep, least caudal peduncle depth 9.7-14.0 (11.0)% SL (36.2-50.3 (42.2)% HL; 105.1-172 (124.3)% caudal peduncle length), greatest caudal peduncle depth 12.0-14.6 (13.2)% SL (44.7-56.0 (50.5)% HL); caudal peduncle length 7.8-10.3 (8.9)% SL (27.5-39.4 (34.1)% HL); caudal-fin length 18.8-25.1 (21.2)% SL. Vertebrae 39-42, with 11 precaudal and 28-31 caudal.

Scales on head, pectoral region, and those scattered posteriorly on ocular side slightly rough and having columnar tubercles in larger adults. Small spines present in small adults.

In life, blind side of adults translucent to white with glossy highlights along edges of myotomes (Amaoka et al., 1983, Fig. 220 as L. bilineata; Amaoka et al., 1995, Fig. 524 as Pleuronectes bilineatus). When preserved, blind side uniform creamy white to yellow-brown. Ocular side brownish with faint yellow highlights around small dark spots near bases of dorsal and anal fins and at midline (Amaoka et al., 1995, Fig. 524).

Remaining description as for genus. Largest specimen examined 280.3 mm (USNM 77130). Maximum size reported 400 mm SL (Sakamoto, 1984b).

Description of juveniles

No juveniles were examined for this study. Juveniles are not described in available literature.

Description of larvae

No larvae were examined in our study. The following description is based on Okiyama and Takahasi (1976, based on 3 specimens of 4.12-8.59 mm) and Pertseva-Ostroumova (1961; number and size of specimens examined not stated).

Snout-to-anus length 32.8-37.0% SL, decreasing with development according to Pertseva-Ostroumova, 1961 (34.2-32.5% SL); body depth 9.0-17.0% SL, increasing with development; head length 15.0-20.0% SL, increasing with development; orbit length 41.0-28.0% HL, decreases with development. Total myomeres 41-43.

Hatching occurring at relatively large sizes, at 3.95-4.48 mm (Yusa, 1958); flexion beginning by 8.9 mm (Fig. B of Okiyama, 1988); postflexion size between 10.6 mm and 15.3 mm (Figs. C and D of Okiyama, 1988); transformation occurring at sizes larger than 15.3 mm (Fig. D of Okiyama, 1988).

Preanal pigment (Fig. 20, based on Okiyama, 1988) present initially along lower jaw, extending ventrally along gut to anus. By flexion, melanophores appearing on pectoral-fin rays; pectoral-fin rays and base not pigmented in all earlier developmental stages; pectoral fin pigmented more heavily than L. polyxystra n. sp. by later stages (Pertseva-Ostroumova, 1961).

Postanal pigment of preflexion larvae in prominent patches of melanophores along finfold edges, three along dorsal and two along anal finfold, becoming more dispersed and less prominent with development; three distinct pigment patches along the dorsal midline, anterior patch begins at about myomere 20-22, posterior patch begins at about 33-36, and caudal patch, which aligns with a ventral patch to form a caudal bar, begins in the range of myomeres 40-44; these dorsal midline patches align vertically with the two posteriormost dorsal and both ventral finfold patches; a series of melanophores may occur along the postanal ventral midline (additional descriptions cannot be found in the literature); additional pigment spots above and below the notochord tip.

Distribution

Lepidopsetta mochigarei ranges in the western Pacific Ocean from the east coast of Korea, Yoglin Bay, in the Sea of Japan (Kim and Youn, 1994) to Iturup Island of the Kuril Islands in the southern Okhotsk Sea (Nikiforov et al., 1983), and to the west coast of northern Japan. Larvae have been collected in March along the coast of northern Honshu in the Sea of Japan (Okiyama, 1988).

Habitat

Adults have been collected over the continental shelf; spawning may occur in depths of about 120 m (Okada, 1955).

Life history

Fecundity ranges from 510,000 to 550,000 eggs (Okada, 1955). Spawned eggs are demersal and adherent, about 0.87-0.95 mm in diameter (Yusa, 1958; Pertseva-Ostroumova, 1961); oocytes at maturation stage IV were 0.4400.655 mm in diameter (Nikiforov et al., 1983). Spawning occurs from December to June in waters around Hokkaido and the southern Kuril Islands (Minami, 1995). Embryonic and larval development to about 5.0 mm has been described by Yusa (1958).

Etymology

The species name mochigarei is derived from the Japanese, meaning “rice-cake flounder” (Snyder, 1911), probably a reference to the bright white blind side similar to white rice-cake common in Japan.(12)

Comments

Masutomi and Hamada (1966) described a fossil of L. mochigarei. It was subsequently described as the extinct Chibapsetta dolichurostyli by Sakamoto and Uyeno (1988).

Because early life history material was not available for examination, descriptions were based on the available literature (Yusa, 1957, 1958; Pertseva-Ostroumova, 1961; Okiyama and Takahashi, 1976; Okiyama, 1988). Several potentially important characters could not be gleaned from the available sources: an anterior pigment spot along the dorsal midline, important in distinguishing L. bilineata from L. polyxystra n. sp. (Figs. 17C, 17D), is not present in preflexion larvae (Figs. 20A and 20B). Because the sources were different, we needed verification that these figures constitute a series. Although present, the extent of the postanal ventral melanophores, important in distinguishing Lepidopsetta from Psettichthys in particular as well as from other pleuronectids, is not known. Additional data for morphological characters were also needed for all developmental stages. Larvae of L. mochigarei appear more similar to those of L. bilineata than L. polyxystra n. sp., but they are not sympatric with L. bilineata. Based on the present descriptions, larvae of L. mochigarei can be distinguished from those of L. polyxystra n. sp. by the distinctive vertically aligned dorsal midline and finfold pigment.

Lepidopsetta polyxystra n. sp. Northern rock sole Figs. 1-5, 8, 11, 13, 21-24; Tables 2-11, 13-14

[Figures 21-24 ILLUSTRATION OMITTED]

Lepidopsetta bilineata var. umbrosa Jordan and Evermann, 1898:2643 (in part, Fig. 928).

Lepidopsetta bilineata bilineata Taranetz, 1937:144 (in part, new combination, keys).

Holotype

UW 014826, 1(169.7 mm), Alaska, Aleutian Is., Amchitka I., Constantine Harbor, 19-42 m, 19 August 1955.

Material examined

A total of 843 adults and juveniles, including the holotype listed above, was examined. Sixty larvae were examined.

Paratypes

Adults 219 specimens, 26-427.5 mm. Okhotsk Sea: UW 042341 (orig. KIE 1147), 2(105.5-109.5 mm), western Kamchatka, 51 [degrees] 57’N, 156 [degrees] 25’E; UW 042342 (orig. KIE 1149), 3(144.5-195 mm), western Kamchatka, 52 [degrees] 02’N, 155 [degrees] 58’E, 6 June 1992; UW 042343 (orig. KIE 1151), 10(186-247 mm), 52 [degrees] 29’N, 153 [degrees] 40’E, 24 June 1992; KIE 1148, 1(89.5 mm), 51 [degrees] 43’N, 156 [degrees] 04’E, 5 June 1992; Southeast Kamchatka: CAS-SU 122549, 1(112.3 mm), Kamchatka, off Petropavlovsk, 18 June 1906; Bering Sea: KIE 1415, 1(289.5 mm), Bering I., SW of Cape Monati, 54 [degrees] 37’N, 166 [degrees] 31’E, 27 April 1996; KIE 1417, 1(258.5 mm), Bering I., SW of Cape Monati, 54 [degrees] 39’N, 166 [degrees] 29’E, 8 May 1996; KIE 1414, 1(260.5 mm), SW of Cape Monati, 54 [degrees] 40’N, 166 [degrees ]29’E; KIE 1268, 1(338.6 mm), NW of Bering I., 55 [degrees] 34’N, 164 [degrees] 55’E, 7 January 1994; KIE 1418, 1(289 mm), Bering I., SE of Cape Monati, 54.6667 [degrees] N, 166.8333 [degrees] E, 9 May 1996; KIE 1416, 1(261 mm), Kamchatka, SW of Cape Northwestern, 55.2833 [degrees] N, 165.5333 [degrees] E, 4 May 1996; KIE 1150, 1(238.5 mm), Kamchatka, 56.2333 [degrees] N, 154.3 [degrees] E, 18 June 1992; ABL 93-37, 1(193 mm), St. Paul I., Pribilof Is.; ABL 60-15, 1(265 mm), N of Pribilof Is., 58 [degrees] 33’N, 170 [degrees] 12’W, 17 July 1960; UW 041694, 3(150-177), Aleutian Is., NE of Umnak I., 53.44 [degrees] N, 168.4944 [degrees] W; SIO 94-164, 9(103-190 mm), Aleutian Is., off Unalaska I., 54 [degrees] 15.1’N, 165 [degrees] 57.6’W, 4 June 94; UW 041199, 1(267 mm), 60.6667 [degrees] N, 169.65 [degrees] W, 16 July 1979; UW 041697, 1(245 mm), Bering Sea, N of Unimak I., 55.0213 [degrees] N, 164.6009 [degrees] W; CAS 45562, 1(240.8 mm), WNW of Pribilof Is., NE end of Zhenchug Canyon, 58 [degrees] 15.8667’N, 174 [degrees] 0.3’W, 21 June 79; CAS 45567, 1(272.1 mm), SSW of Pribilof Is., 55 [degrees] 38.8667’N, 168 [degrees] 39.9833’W, 16 June 79; CAS 46676, 1(87.7 mm), Pribilof Is., St. George I., 56 [degrees] 35’N, 169 [degrees] 36’W, 27 July 1977; CAS 47531, 5(181.4-199.4 mm), Aleutian Is., 51 [degrees] 43.9’N, 177 [degrees] 57.8833’W, 10 August 1980; CAS-SU 105731, 5(132.6-182.9 mm), Aleutian Is., Umnak I., off Nikolski, 31 July 1896; USNM 130725, 3(287-310 mm), 40 mi above Port Moller, 8 May 1941; USNM 060905, 1(316.5 mm), Karluk, TT2204, RV Albatross; UW 001378, 1(175.5 mm), Unalaska I., 1930; UW 003656, 1(100.5 mm), Alaska Peninsula, Cold Bay, 13 June 1932; UW 013667, 1(208.1 mm), Unimak I., False Pass, 1 July 1957; UW 016609, 1(191 mm), Pribilof Is., St. George I.; UW 025736, 3(28-45.7 mm), Port Moller; UW 016600, 3(83.9-130.9 mm), 58.5833 [degrees] N, 168.1 [degrees] W, 4 July 1949; UW 003100, 3(86.7-164.6 mm), Captains Bay, Unalaska I.; UW 10650, 2(81.6-112.8 mm), off Nunivak I., 59 [degrees] 36’N, 167 [degrees] 56’W; UW 14831, 1(83 mm), off Nunivak I., 59.82 [degrees] N, 169.13 [degrees] W, 4 July 1949; UW 25760, 1(193 mm), 57.65 [degrees] N, 167.65 [degrees] W; UW 25731, 1(178.9 mm), 57.6667 [degrees] N, 167.2 [degrees] W, 27 April 1987; UBC 65-39, 1(175 mm), Izembeck Bay, 31 August 1964; UBC 62-565, 2(198-229 mm), 120 km E of St. George I.; UBC 65-730, 2(220-245 mm), 57 [degrees] 45’N, 164 [degrees] 45’W, 16 August 1965; UBC 65-714, 1(160 mm), 57 [degrees] 45’N, 168 [degrees] 45’W, 1 August 1965; UBC 65-729, 1(150 mm), 55 [degrees] 45’N, 162 [degrees] 45’W, 14 August 1965; Gulf of Alaska: ABL 72-74, 6(26-59.3 mm), SE shore of Favorite Channel, 10 November 1972; ABL Uncat, 1(195 mm), Unalaska I., 18 June 1958; 86ABL 62-202, 3(83.5-88 mm), Auke Bay, ca. 17.6 km NW of Juneau, 23 May 1962; ABL 62-199, 1(52.3 mm), 19.2 km NW of Juneau; ABL 64-996, 2(65.6-69.5 mm), Auke Bay, ca. 17.6 km NW of Juneau, 30 April 1964; ABL 67-32, 1(265 mm), Baranof I., Katlian Bay, 19 April 1967; OS 17158, 20(227-315 mm), 55.3323 [degrees] N, 161.3186 [degrees] W, 32 m, 3 June 1996; OS 17157, 3(230-262 mm), Aleutian Is., 51.88373 [degrees] N 179.7202 [degrees] E, 95 m, 16 July 1997; OS 3497, 1(213 mm), False Pass, Ikatan Bay, 19 June 1956; OS out of 6447, 2(166.1-170.2 mm), 57 [degrees] 10’N, 151 [degrees] 40.5’W, 77 m, 20 September 1978; OS 2157, 1(92 mm), Kodiak I., Shelikof St., mouth of Sturgeon R., 9-10 September 1954; OS 2169, 1(64 mm), Moffet Pt. to Black Hill, 28 May 1957; OS 2173, 1(63.5 mm), Moffet Pt. to Black Hill, 3 June 1957; OS 3979, 1(94.6 mm), Little Port Walter, 8 August 1964; SIO 69-748, 3(51-195 mm) of 7(51-195 mm), Afognak I., Kitoi Bay, 16-21 April 1968; SIO 76-299, 1(270 mm), Kodiak shelf, 56 [degrees] 43’N, 153 [degrees] 22’W, 25 June 1976; SIO 72-227, 1(235 mm), 40 km NW of Juneau, Lynn Canal, 16 February 1963; USNM 130734, 1(254.5 mm), Ivanof Bay, 3 April 1941; USNM 27602,1(261.8 mm), Kodiak I., 12 July 1880; USNM 27942, 1(260.2 mm), Cook Inlet, Port Chatham; USNM 116332, 2(293-295 mm), Canoe Bay, 17 September-15 October 1940; UW 003854, 1(355 mm), Alitak Bay, Kodiak I., 9 June 1932; UW 040498, 1(195 mm), Prince William Sound, 1989; UW 025740, 1(68.4 mm), Ugak Bay, Kodiak I.; UW 044029, 1(276 mm), 59.5743 [degrees] N, 151.325 [degrees] W, 11 m depth, 26 February 1996; UW 044001, 18(52.7-99 mm) [3(30-40 mm) of 18 used for illustration], Prince William Sound; UBC 59-485, 2(71-116.2 mm), Chief Cove, Spiridon Bay; UBC 65-572, 1(74 mm), Boussole Bay, 29 June 1965; UBC 63-174, 1(290 mm), Prince William Sound, off Cordova; UBC 65-148, 1(205 mm), W of Chirikof I., 3 August 1964. British Columbia: SIO 63-206-64B, 1(97 mm), Vancouver I., Stanley Park at Lumberman’s Arch, 15 March 1963; UBC 56-83, 3(83.5-121.3 mm), Strait of Georgia, off Saturna I.; UBC 62-874, 1(120 mm), off Sooke, 20 June 1962; Puget Sound: OS 3565, 1(224 mm), Puget Sound, 3 February 1943; UW 041078, 2(126-144 mm), Nisqually Reach, 25 February 1949; UW 004411, 1(36 mm), Edmonds, 1 June 1938; UW 000203, 1(45.9 mm), Lopez I., 12 August 1929; UW 041678, 1(67.2 mm), San Juan Islands, East Sound, 11 December 1964; UW 041675, 3(53.7-166.5 mm), Port Townsend Bay, 5 December 1978-5 January 1979; UW 018096, 1(182.4 mm), Bellingham Bay, Chuckanut Bay, May 1964; UW 044014, 29(121-223 mm), 47 [degrees] 19.48’N, 122 [degrees] 33.8’W, 10 May 1996; UW 044007, 9(135-200 mm), Nisqually, 47 [degrees] 9.47’N, 122 [degrees] 40.16’W, 16 May 1996.

Larvae 18 specimens, 3.3-18.6 mm, examined. Bering Sea: UW 083475, 1(3.7 mm), 55 [degrees] 47.0’N, 165 [degrees] 59.0’W, 0-117 m depth, bongo net, 19 April 1991; Gulf of Alaska: UW 083456, 1(3.3 mm), 58 [degrees] 22.0’N, 150 [degrees] 06.0’W, 0-52 m depth, bongo net, 9 April 1991; UW 083479, 1(11.2 mm), 57 [degrees] 19.0’N, 156 [degrees] 09.0’W, 0-100 m depth, bongo net, 4 June 1990; UW 083457, 1(4.2 mm), 56 [degrees] 30.0’N, 157 [degrees] 00.2’W, 0-43 m depth, bongo net, 24 April 1991; UW 083476, 1(5.2 mm), 56 [degrees] 54.3’N, 156 [degrees] 15.3’W, 0-100 m depth, bongo net, 9 May 1992; UW 083461, 1(6.4 mm), 56 [degrees] 46.1’N, 156 [degrees] 47.9’W, 0-47 m depth, bongo net, 24 May 1991; UW 083472, 1(7.2 mm), 56 [degrees] 46.2’N, 156 [degrees] 33.6’W, 0-99 m depth, bongo net (333 mesh), 13 May 1992; UW 083471, 1(8.8 mm), 56 [degrees] 46.2’N, 156 [degrees] 33.6’W, 0-99 m depth, bongo net, 13 May 1992; UW 083478, 2 of 10 (10.0-10.7 mm), 56 [degrees] 56.7’N, 154 [degrees] 43.2’W, 0-42 m depth, bongo net, 5 June 1990; UW 083481, 2(13.6-14.2 mm), 56 [degrees] 54.0’N, 156 [degrees] 29.5’W, 0-84 m depth, bongo net, 3 June 1990; UW 083477, 1(5.7 mm), 55 [degrees] 57.0’N, 158 [degrees] 33.1’W, 0-100 m depth, bongo net, 21 May 1991; UW 083480, 2(12.4-12.6 mm), 55 [degrees] 39.1’N, 156 [degrees] 3.4’W, 0-100 m depth, bongo net, 31 May 1990; UW 083451, 1(17.8 mm), 55 [degrees] 18.1’N, 160 [degrees] 12.0’W, 0-115 m depth, Methot net, 24 July 1991; UW 083450, 1(15.3 mm), 54 [degrees] 39.2’N, 159 [degrees] 23.0’W, 0-54 m depth, Methot net, 29 July 1991; UW 083452, 1(18.6 mm), 54 [degrees] 36.2’N, 162 [degrees] 16.2’W, 0-87 m depth, Methot net, 30 July 1991.

Other material examined

Adults 629 adults and juveniles. Bering Sea: UW 044023, 6(173-292 mm), 60.6869 [degrees] N, 175.4609 [degrees] W, 11 July 1995; UW 044020, 2(331-368.5 mm), 58.45 [degrees] N, 174.33 [degrees] W; UW 044003, 2(226-249 mm), 58.12 [degrees] N, 168.43 [degrees] W; UW 041699, 1(108 mm), 58.66 [degrees] N, 159.47 [degrees] W; Gulf of Alaska: UW 044015, 307(18.2-190 mm), Kachemak Bay; UW 044016, 1(37.5 mm), Kachemak Bay; UW 044018, 54(26-90 mm), Kachemak Bay; SIO 76-300, 3(161-200 mm) of 28(145-280 mm), Kodiak shelf, 57 [degrees] 40’N, 150 [degrees] 37’W, 15 March 1963; ABL 68-505, 2(128-140 mm) of 15(15-140 mm) examined, Glacier Bay, 8 August 1968; UW 025735, 2(35.8-61.8 mm), “6/10/83, Sta. 14, Haul 52”; UW 04321, 1(230 mm), Fishing Vessel Sulak, Cruise 436, Robin Scheid; UW 002688, 83(56-107 mm), Shumagin Is., Baralof Bay, 27 June 1931; UW 002032, 30(75-137 mm), Yakutat Bay, 21 June 1932. UW 041674, 11(67.5-139 mm), Alexander Archipelago, off Wrangell I., 1 December 1931; UW 003841, 13(72-99 mm), Yakutat Bay, 21 June 1932; UW 004004, 7(77-100 mm), Yakutat Bay, 21 June 1932; UW 005047, 6(97-120 mm), Alitak Bay, 9 June 1932; UW 041681, 1(119.5 mm), Prince William Sound, 1989; UW 002069, 8(106-157.3 mm), Cold Bay, 6-26 May 1932; UW 022413, 5 cleared and stained of 14(53-110 mm), Adak I., Aleutian Is.; UW 040265, 16(159-350 mm), 54.03899 [degrees] N, 165.8153 [degrees] W; UW 003913, 5(113.4-167 mm), Prince William Sound, Orca Inlet, 2 April 1935; UW 044012, 6(275-384 mm), 56.5 [degrees] N, 153.5 [degrees] W, Bonaduhr; UW 020658, 7(56.1-135.7 mm), Kodiak I., Ugak Bay; UW 025763, 1(139.7 mm), off Amchitka I.; UW 040263, 3(214-255 mm), 54 [degrees] N, 160.74 [degrees] W. British Columbia: UBC out of 61-609, 3(44.8-56.6 mm), Bute Inlet; UBC out of 60-416, 6(181-230 mm), Queen Charlotte Is., Gillat Arm; UBC out of 62-93, 7(75-111 mm), Nass Bay; UBC out of 61-674, 7(64-81 mm), Dean Channel, off Nescall Bay. Washington, Oregon, and California: UW 041673, 5(68-110.5 mm), San Juan Is., East Sound, 3 March 1937; UW 047270, 6(60-240 mm), Puget Sound, Port Townsend Bay; CAS 19305, 1(246.5 mm), San Francisco (see comments).

Larvae 42 specimens (4.2-20.0 mm) examined: Bering Sea: UW 083446, 1(10.8 mm), 57 [degrees] 30.0’N, 169 [degrees] 30.0’W, depth unknown, bongo net, 26 July 1971; UW 083447, 1(4.5 mm), 54 [degrees] 01.3’N, 166 [degrees] 33.9’W, 0-100 m depth, bongo net, 25 April 1993; Western Gulf of Alaska: UW 083448, 1(15.1 mm), 55 [degrees] 17.7’N, 160 [degrees] 11.7’W, 0-115 m depth, Methot net, 24 July 1991; UW 083449, 1(15.3 mm), 54 [degrees] 39.2’N, 159 [degrees] 23.0’W, 0-54 m depth, Methot net, 29 July 1991; UW 071740, 1(12.0 mm; 11.0 mm as measured in this study), 55 [degrees] 38.6’N, 153 [degrees] 30.1’W, 0-102 m depth, bongo net, 31 May 1990; UW 071752, 1(10.0 mm; 9.4 mm as measured in this study), 55 [degrees] 31.6’N, 156 [degrees] 16.3’W, 0-100 m depth, bongo net, 31 May 1990; UW 072106, 1 of 2(17.1 mm), 56 [degrees] 46.2’N, 156 [degrees] 33.6’W, 0-100 m depth, bongo net, 3 June 1990; Gulf of Alaska: UW 069346, 1(4.3 mm) of 9, 58 [degrees] 22.0’N, 150 [degrees] 12.8’W, 0-47 m depth, bongo net, 30 March 1978; UW 083453, 1(12.7 mm), 56 [degrees] 41.9’N, 154 [degrees] 33.2’W, 0-16 m depth, bongo net, 24 June 1978; UW 083454, 1(16.3 mm), 57 [degrees] 19.5’N, 152 [degrees] 23.9’W, 0-48 m depth, bongo net, 24 June 1978; UW 083455, 1(7.4 mm), 56 [degrees] 23.7’N, 155 [degrees] 45.0’W, 0-79 m depth, bongo net, 15 May 1979; UW 063735, 1 of 12 (11.8 mm), 56 [degrees] 35.1’N, 153 [degrees] 43.6’W, 0-70 m depth, bongo net, 25 May 1985; UW 072459, 1 of 10 (9.4 mm), 57 [degrees] 10.7’N, 155 [degrees] 12.9’W, 0-100 m depth, bongo net, 5 June 1990; UW 072497, 6 of 10 (7.4-12.6 mm), 56 [degrees] 56.7’N, 154 [degrees] 43.2’W, 0-42 m depth, bongo net, 5 June 1990; UW 072502, 4 of 9 (7.3-10.3 mm), 56 [degrees] 49.4’N, 154 [degrees] 30.3’W, 0-60 m depth, bongo net, 5 June 1990; UW 072511, 1(12.2 mm), 57 [degrees] 28.6’N, 154 [degrees] 42.4’W, 0-60 m depth, bongo net, 5 June 1990; UW 072546, 1(8.1 mm), 57 [degrees] 40.6’N, 155 [degrees] 10.5’W, 0-287 m depth, bongo net, 6 June 1990; UW 083458 1(6.8 mm), 57 [degrees] 15.6’N, 155 [degrees] 55.4’W, depth unknown, Mocness net, 14 May 1991; UW 083460, 1(6.4 mm), 56 [degrees] 16.4’N, 158 [degrees] 04.9’W, 0-64 m depth, bongo net, 22 May 1991; UW 083463, 1(4.2 mm), 57 [degrees] 52.3’N, 155 [degrees] 0.1’W, 0-82 m depth, bongo net, 7 April 1992; UW 083464, 1(5.0 mm), 57 [degrees] 42.2’N, 154 [degrees] 47.3’W, 0-219 m depth, bongo net, 7 April 1992; UW 083465 1(5.5 mm), 57 [degrees] 16.5’N, 155 [degrees] 55.0’W, 0-100 m depth, bongo net, 8 May 1992; UW 083466, 1(5.8 mm), 56 [degrees] 54.3’N, 156 [degrees] 15.3’W, 0-100 m depth, bongo net, 9 May 1992; UW 083467 3(4.6-6.0 mm), 56 [degrees] 51.8’N, 156 [degrees] 21.4’W, 0-99 m depth, bongo net, 9 May 1992; UW 083468, 1(6.1 mm), 56 [degrees] 53.3’N, 156 [degrees] 23.9’W, 0-102 m depth, bongo net, 9 May 1992; UW 083469, 1(8.5 mm), 56 [degrees] 42.2’N, 156 [degrees] 32.3’W, 0-100 m depth, bongo net, 13 May 1992; UW 083470, 1(6.7 mm), 56 [degrees] 44.4’N, 156 [degrees] 28.7’W, 0-102 m depth, bongo net, 13 May 1992; UW 083473 1(7.6 mm), 56 [degrees] 09.4’N, 157 [degrees] 14.3’W, 0-101 m depth, bongo net, 22 May 1992; UW 083474, 1(7.5 mm), 56 [degrees] 40.6’N, 155 [degrees] 10.4’W, 0-59 m depth, bongo net, 15 May 1992; UW 083482, 2(19.0-20.0 mm), 47 [degrees] 34.15’N, 122 [degrees] 32.30’W, 0 m depth, dip net, 9 May 1990.

Diagnosis

Lepidopsetta polyxystra is a species of Lepidopsetta with the following combination of characters in adults: total gill rakers on first arch 9-14, on upper arch 2-6; total gill rakers on second arch 8-14; supraorbital pores 1-3, rarely 4-7; preopercular pores 5-8; lateral-line pores 76-100; sum of scales above and below lateral line 66-96; interorbital wide; blind side coloration in life creamy, without glossy highlights along margins of myotomes.

Larvae are distinguished from other species of Lepidopsetta by the following characters: body slender; snout-to-anus length long; larvae undergo hatching, flexion, and transformation at comparatively larger sizes; preflexion pigment pattern with light pigmentation along finfolds, limited mainly to the ventral finfold, and two pigment spots along the dorsal midline (the posteriormost aligning with a ventral patch forming a bar), a series of small melanophores extending from just posterior to the anus to just beyond the postanal bar (about 2/3 body); and flexion pigment pattern with a single bar located on the postanal body; pectoral-fin rays unpigmented.

Description of adults

Body ovate, greatest depth 41.2-58.9 (49.0)% SL, scales above lateral line 27-40, scales below lateral line 38-59; head relatively acute, length 21.8-31.5 (27.2)% SL; dorsal margin of head at dorsal-fin origin slightly concave, snout length 3.1-5.7 (4.4)% SL (12.1-20.1 (16.0)% HL); ocular-side maxilla length 20.4-31.5 (26.3)% HL; blind-side maxilla length 22.2-35.7 (28.7)% HL; ocular-side mandible length 34.4-50.0 (40.7)% HL; teeth 4-5 on ocular-side premaxilla, 23-24 on blind-side premaxilla and 8-9 on ocular-side dentary, 20-24 on blind-side dentary; gill rakers of first arch relatively slender, 9-14 total, 3-6 on upper arch, 6-9 on lower arch; gill rakers of second arch 8-14 total with 0-4 on upper and 7-13 on lower arch; dorsal orbit length often with posteriorly elongate rim and much longer than eye length, orbit length 22.2-32.9 (28.1)% HL, dorsal eye length 18.5-32.9 (23.0)% HL; ventral eye length 17.2-31.1 (24.2)% SL; interorbital wide, up to 5 scales at narrowest portion, 2.5-6.6 (4.4)% HL; cheek with 7-12 scales, length 26.7-38.5 (32.5)% HL, depth 13.6-22.7 (18.7)% HL; preopercular pores 5-8; ocular-side suborbital pores 13-29; blind-side suborbital pores 8-14; lateral line pores 76-100, lateral-line arch length 26.1-64.5 (52.3)% HL, its depth 26.3-38.2 (31.7)% its length; both anterior and posterior supratemporal branches relatively long, anterior pores 3-15, posterior pores 10-35; supraorbital canal short, reaching only to the posterior rim of the dorsal orbit at about the insertion of dorsal-fin rays 5-6, pores 1-3, rarely 4-7; ocular-side pectoral-fin length 10.7-18.2 (14.7)% SL (38.3-70.4 (53.9)% HL); blind-side pectoral-fin length 6.6-12.7 (10.2)% SL (24.2-49.6 (37.4)% HL); ocular-side pelvic-fin length 7.8-12.2 (9.9)% SL (27.6-45.9 (36.6)% HL); dorsal fin with 64-83 rays, height 10.2-16.6 (13.5)% SL, in specimens with 69-80 rays supported by 68-78 pterygiophores, typically 9 or rarely 8 anterior to first neural spine; anal fin with 49-64 rays, in specimens with 55-61 rays supported by 53-63 pterygiophores; caudal peduncle relatively slender, least depth 9.4-13.1 (10.9)% SL (32.2-48.8 (40.2)% HL, 101.2-144.2 (123)% caudal peduncle length), greatest depth 10.1-16.9 (12.3)% SL (38.1-57.7 (45.2)% HL); caudal peduncle length 7.0-10.1 (8.9)% SL (27.6-42.0 (32.7)% HL); caudal-fin length 17.8-29.1 (22.7)% SL. Vertebrae 39-41, with 10-11 precaudal and 29-30 caudal.

Scales around head and those scattered posteriorly on ocular side very rough, with columnar tubercles in large adults; strong spines in small adults and juveniles. Urogenital flap unpigmented or lightly pigmented in 23-80 mm juveniles.

In life, blind side in moderate to large adults creamy white, skin opaque (Amaoka et al., 1995, Fig. 525); in juveniles, blind side nearly completely translucent, with small, white glossy areas on head. When preserved, blind side of all individuals uniform creamy white to yellow-brown. Ocular side slightly less green than that of congeneric in Puget Sound, often with faint yellow highlights around darker spots near bases of dorsal and anal fins and at midline.

Remaining description as for genus. Largest specimen examined 340 mm (427.5 mm TL) (UW 040265). Maximum size reported 588 mm (Fadeev, 1965) to 690 mm TL.(6)

Description of juveniles

Individuals of about 20 mm collected in water column (one 19.0-mm individual examined, UW 083482, not completely transformed; see comments); lateral line and pectoral-fin rays undeveloped; body pigmentation increasing throughout; bars and patches of postflexion larval pigmentation pattern visible; urogenital papilla light or speckled with an unpigmented tip. Eye length smaller, mouth larger, body deeper, gill-raker counts on lower arch higher, distance from pelvic-fin origin to anal-fin origin shorter, expanded anteriormost anal-fin pterygiophore less developed (not protruding beyond body wall in our material) than in similar-size L. bilineata.

By 35 mm, many individuals collected near bottom. Postsettlement juveniles (Fig. 22B) as developed as similar-size L. bilineata: increased body pigmentation giving juveniles a darker appearance (obscuring urogenital papillae pigment), rays of paired and median fins formed, lateral line formed, supraorbital pores present, expanded anteriormost anal-fin pterygiophore strongly developed.

Description of larvae

Snout-to-anus length 32.1-39.3% SL, remaining constant during development; body depth 3.8-28.5% SL, increasing with development, sharply increasing after flexion; head length 11.6-26.7% SL, increasing with development; snout length 22.8-24.7% HL, remaining constant during development; orbit length 51.8-20.9% HL, decreasing with development (Table 2). Total myomeres 38-42.

Larvae hatching at greater than 3.0 mm (3.6-4.0 mm, Pertseva-Ostroumova, 1961); yolk absorbed by 3.3-4.2 mm. Preflexion larvae ranging in size from 4.2 to 6.2 mm; flexion larvae, 6.2-12.6 mm; postflexion larvae, 12.2-18.6 mm. Transformation occurring at sizes as small as 15.0 mm; postsettlement stage usually not complete by 20.0 mm (Table 13).

Head pigment present initially along lower jaw and underside of chin (Fig. 23); increasing with development to snout, upper jaw, and isthmus. Pigment ventrally along gut and dorsally on anus; by flexion a distinct patch of melanophores along posterior edge of gut; pigment increasing laterally with development.

Postanal pigment light along the anal finfold, melanophores absent on dorsal finfold (Fig. 23); two pigment patches along the dorsal midline, anterior patch beginning about midbody at myomere 18-20, posterior patch beginning about myomere 30, posterior patch aligning with a ventral patch forming a postanal bar; series of melanophores along the ventral body midline beginning just posterior to the anus, extending to just beyond the postanal bar (about 2/3 body length); a few melanophores on caudal peduncle and above and below the notochord tip.

Distribution (Figs. 8, 11, 13, 24)

Ranging from the northern coast of Hokkaido throughout the Kuril Islands and the Okhotsk Sea of the western North Pacific, through the Bering Sea, from the Gulf of Anadyr(13) to off St. Lawrence Island (Allen and Smith, 1988), to Puget Sound, Washington, in the eastern North Pacific. Large concentrations have been reported from the west and southeast of the Kamchatka Peninsula, where it once composed 90% of the fisheries catches of pleuronectids (Shubnikov and Lisovenko, 1964), and from the southeastern Bering Sea. Minami and Nakamura (1978) also reported a single specimen of L. bilineata (=L. polyxystra n. sp. based on distribution) among many L. mochigarei from Wakasa Bay, Japan (ca. 35.7 [degrees] N, 135.5 [degrees] E); this specimen could not be located.(14) Larvae have been collected from Puget Sound, the east coast and northern tip of Vancouver Island, Hecate Strait, and north along the coasts of Alaska into the Bering Sea. According to Okiyama (1988), larvae occur along the coast of the Kuril Islands, the coast of Kamchatka, and into the Bering Sea.

The recorded locality of one adult (CAS 19305) from San Francisco Bay, over 1200 km south of the nearest verified capture in Puget Sound, is questionable. Although no evidence from catalog records(8) indicates that the specimen had been misplaced, the specimen was collected in 1888 and has an original catalog number from the Indiana University collection. Its morphology and meristics are distinctively that of more northerly populations.

Habitat

Lepidopsetta collected in the eastern Bering Sea are most commonly associated with sand, and least with “sand and mud,” when compared with all other measured substrate types in the Bering Sea, including combinations of sand, gravel, and mud.(15) The maximum depth of collection was 246 m.(16) During the 22-year ichthyoplankton sampling period (all previously unidentified pleuronectid larvae were re-examined, thus providing the only historic distributional records for larvae of L. polyxystra n. sp.), larvae were common in spring surveys conducted in the Gulf of Alaska and Bering Sea, when they first appear in March collections; largest catches have been taken in May. Highest densities of larvae occur in the Bering Sea (Fig. 24), compared with the western Gulf of Alaska.

Life history

Lepidopsetta collected from the eastern Bering Sea and Sakhalin Island, presumed to be L. polyxystra n. sp., fed primarily on polychaetes and other marine worms; molluscs (Skalkin, 1959, 1963; Lang et al., 1995) or fishes, primarily Ammodytes hexapterus in depths [is less than] 50 m (Lang et al., 1995), were a second significant component of the diet. Feeding took place primarily from May to September, falling to low levels during the winter (Shubnikov and Lisovenko, 1964) when spawning occurs (Fadeev, 1965). Spawning occurs in areas with good water circulation over hard bottoms of sand with gravel and peaks from early March to mid-April in waters off Kamchatka, possibly occurring as late as mid-June in the western Pacific (Pertseva-Ostroumova, 1961). Fecundity ranges from 151,700 to 404,200 eggs (Fadeev, 1965). Life history including spawning and development was discussed by Pertseva-Ostroumova (1961, as L. bilineata bilineata) and larval development was described by Okiyama (1988, as L. bilineata).

The Washington Department of Fisheries reported Lepidopsetta from late December through early March in sandy gravel of upper intertidal beaches at several sites in central and southern Puget Sound (Penttila, 1995). In January 1991, one beach site on the south shore of Dana Passage (the center of a relatively large area of documented Lepidopsetta spawning) was the source of six batches of field-collected spawned eggs (each about 1 mm in diameter) that were subsequently reared through hatching. We identified the reared larvae as L. polyxystra n. sp. Egg batches ranged in size from 40 to several hundred eggs and were incubated over about 14 days with ambient central Puget Sound water (about 9.0 [degrees] [C.sup.3]). Larvae hatched between 4 and 5 mm and had yellow pigmentation associated with the melanophores. For additional early life history information, see generic account.

The maximum recorded age for female L. polyxystra n. sp. captured in the Bering Sea is 18 yr at 49 cm FL, and for males is 17 yr at 40 cm FL (Alton and Sample(17)). The gills of adults from the Gulf of Alaska have been found to be heavily infested with copepod parasites Nectobrachia indivisa, and Naobranchia occidentalis, previously recorded in Lepidopsetta by Kabata (1988). For both parasites, L. polyxystra n. sp. was significantly more heavily infested than L. bilineata.(7) The parasites Acanthochondria vancouverensis and Haemobaphes sp. were also recorded.

Etymology

The specific name polyxystra is derived from the Greek poly, meaning many, and xystrus, meaning raker, referring to the gill rakers being more numerous than those of congeneric species.

Comments

Under the name of L. bilineata, L. polyxystra n. sp. has been the subject of many studies. Work on specimens collected north of the extreme southeastern Bering Sea and west of the eastern Aleutian Islands in the western North Pacific may be presumed to be based on L. polyxystra n. sp. However, all studies conducted on specimens from the Gulf of Alaska into Puget Sound, regions of extensive overlap in the distributions of L. polyxystra n. sp. and L. bilineata, should be considered applicable at the generic level only, unless voucher specimens were collected and can be verified.

According to Pertseva-Ostroumova (1961), juveniles as small as 20 mm SL have been collected on the bottom. However, the species exhibits much plasticity in settling size, with specimens much greater than 20 mm SL routinely collected in plankton nets.

A widely published illustration identified as L. bilineata is based on a specimen of L. polyxystra n. sp. taken off Kodiak Island, Alaska (“St. Paul, Kodiak,” USNM 27602), now badly damaged. First published in Goode (1884), it was subsequently reprinted in Jordan and Goss (1889), Jordan and Evermann (1900), Jordan and Starks (1906), and Evermann and Goldsborough (1907) among other publications.

Acknowledgments

Beverly Vinter first noticed an unidentified pleuronectid in our ichthyoplankton samples almost 15 years ago. With support from A. W. Kendall Jr., she persisted over the years in her belief that these larvae represented a new species of pleuronectid. We thank her for her insight as well as her identifications, illustrations, measurements, and numerous examinations of larvae. Early in our study, A. W. Kendall Jr. arranged for the collection of adults, the rearing of larvae (with D. Misitano, formerly of NWFSC), and electrophoretic analysis of samples (with F. Utter formerly of NWFSC). Helen Mulligan (first working at AFSC and later on contract to Humboldt State University) made several significant contributions to the study including a preliminary analysis of the geographic distribution of larval and adult Lepidopsetta that provided evidence supporting a taxonomic basis for the observed differences in larval shape and structure. Maryjane Cleveland, a domestic fisheries observer, provided detailed notes of her observations of Lepidopsetta, sparking independent early collections of adult samples. Other samples and data have been provided over the years by the AFSC’s RACE Division (W. Flerx, G. R. Hoff, R. MacIntosh, G. Walters, and K. Weinberg), Resource Ecology and Fisheries Management Division’s Observer Program (S. Barbeaux, M. Brown, S. Corey, K. Kruse, M. Loefflad, R. Narita, N. Raring, and K. Scott), and by the Alaska Department of Fish and Game (H. Sanborn and D. Urban). Biologists of the Washington Department of Fish and Wildlife, including W. A. Palsson, R. Pacunski, and G. Lippert, provided comparative photographs, specimens, and data for the Puget Sound. Jeff Marliave (Vancouver Public Aquarium) also provided Canadian larval material. Data on distribution and abundance of larvae and juveniles have been provided by H. G. Moser (SWFSC), W. Shaw (DFO, PBS), B. L. Norcross, A. A. Abookire, and B. Holladay (UA, Fairbanks), and D. Penttila (WDF). Several scientists at AFSC assisted throughout this study: D. M. Blood and M. S. Busby (collections at sea and technical assistance), L. Ciannelli (measurements), K. Mier (statistical analyses), S. Picquelle (mapping software and CPUE calculations), R. C. Harrison (translations), and W. Rugen (database support). For discussions on pleuronectid characters, osteology, and phylogeny, we thank J. A. Cooper and P. Berendzen. The following curators and collection managers generously provided loans and information on specimens under their care: D. Catania (CAS), W. G. Saul (ANSP), B. A. Brown (AMNH), R. F. Feeney and J. Seigel (LACM), J. D. McPhail, G. Haas, and E. Keeley (UBC), D. F. Markle (OS), B. Sheiko (KIE), T. W. Pietsch and B. K. Urbain (UWFC), S. Jewett and S. J. Raredon (NMNH), R. H. Rosenblatt, H. J. Walker, and C. Klepadlo (SIO), K. Amaoka (HUMZ), G. Shinohara (NSMT), I. Kinoshita (FRSKU), T. Minami (Japan Sea National Fisheries Institute), and B. L. Wing (ABL). We thank the following individuals for their reviews of the manuscript: T. W. Pietsch, W. Aron, G. R. Hoff, M. S. Busby, A. W. Kendall Jr., C. W. Mecklenburg, E. S. Brown, and H. H. Zenger.

(1) Kendall, A. W., Jr., and A. C. Matarese. 1987. Unpubl. data. Resource Assessment and Conservation Engineering Division, Alaska Fisheries Science Center, Natl. Mar. Fish. Serv., NOAA, 7600 Sand Point Way NE, Seattle, WA 98115.

(2) Sheiko, B. 1997. Personal commun. Kamchatka Institute of Ecology, Partizanskaya 6, Petropavlovsk-Kamchatsky 683000, Russia. Present address: Department of Ichthyology, Zoological Institute, Russian Academy of Sciences, Universitetskaynab. 1, St. Petersburg 199034, Russia.

(3) Penttila, D. 1996. Personal commun. Puget Sound Forage Fish Unit, Washington Department of Fish and Wildlife, 1702 Anderson Road, Bay 4, Mt. Vernon, WA 98173.

(4) We do not recognize two other generic nomenclatural changes recommended by Cooper and Chapleau (1998) and have chosen to retain Embassichthys and Atheresthes as separate genera, distinct from Microstomus and Reinhardtius, respectively.

(5) Cooper and Chapleau (1998) changed the specific epithet of Parophrys vetulus to vetula to agree in gender with Parophrys. However, because Girard’s (1854) original description did not specify his treatment of vetulus and because it may also be treated as a noun in apposition, we consider this change an incorrect subsequent spelling and retain P. vetulus.

(6) Resource Assessment and Conservation Engineering (RACE) Division. 1996. Unpubl. data from RACE database. Alaska Fisheries Science Center, Natl. Mar. Fish. Serv., NOAA, 7600 Sand Point Way NE, Seattle, WA 98115.

(7) Zimmermann, M. and R. Harrison. 1998. Personal commun. Resource Assessment and Conservation Engineering Division, Alaska Fisheries Science Center, Natl. Mar. Fish. Serv., NOAA, 7600 Sand Point Way NE, Seattle, WA 98115.

(8) Catania, D. 1996. Personal commun. Ichthyology Department, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118.

(9) Saul, W. 1996. Personal commun. Academy of Natural Sciences, 19th and The Parkway, Philadelphia, PA 19103.

(10) Feeney, R. 1996. Personal commun. Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007.

(11) Jewett, S. 1995. Personal commun. Division of Fishes, Smithsonian Institution, Washington, D.C. 20560.

(17) Alton, M. A., and T. Sample. 1976. Rock sole (family Pleuronectidae). In Demersal fish and shellfish resources of the eastern Bering Sea in the baseline year 1975 (W. T. Pereyra, J. E. Reeves, and R. G. Bakkala), p. 461-474. Northwest and Alaska Fish. Center Proc. Rep., Seattle, WA.

Literature cited

Ahlstrom, E. H., K. Amaoka, D. A. Hensley, H. G. Moser, and B. Y. Sumida. 1984. Pleuronectiformes: Development. In Ontogeny and systematics of fishes (H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall Jr., and S. L. Richardson, eds.), p. 640-670. Am. Soc. Ichthy. Herpet. Spec. Publ. 1.

Allen, M. J., and G. B. Smith. 1988. Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific. U.S. Dep. Commer., NOAA Tech. Rep., NMFS 66, 151 p.

Amaoka, K., K. Nakaya, and M. Yabe. 1995. The fishes of northern Japan. Kitanihon Kaiyo Center Co., Sapporo, Hokkaido, 387 p.

Amaoka, K., K. Nakaya, and H. Yamamoto. 1983. Fishes and marine algae of northern Japan. Kitanihon Kaiyo Center Co., Sapporo, Hokkaido, 371 p.

Ayres, W. O. 1855a. [Description of new fishes from California.] The Pacific 4(18).

1855b. [Description of a new flounder.] Proc. Calif. Acad. Sci. 1:39-40.

Berendzen, P. B. 1998. Phylogenetic analysis of the order Pleuronectiformes using molecular and morphological evidence. M.S. thesis, Univ. Kansas, Lawrence, KS, 61 p.

Blackburn, J. E. 1973. A survey of the abundance, distribution, and factors affecting distribution of ichthyoplankton in Skagit Bay. M.S. thesis, Univ. Washington, Seattle, WA, 136 p.

Busby, M. S., A. C. Matarese, and K. L. Mier. 2000. Annual, seasonal, and diel composition of larval and juvenile fishes collected by dip-net in Clam Bay, Puget Sound, Washington, from 1985 to 1995. U.S. Dep. Commer., NOAA Tech. Memo. NMFS-AFSC-111, 36 p.

Chapleau, F. 1993. Pleuronectiform relationships: a cladistic reassessment. Bull. Mar. Sci. 52(1):516-540.

Charter, S. R., and H. G. Moser. 1996. Pleuronectidae: righteye flounders. In The early stages of fishes in the California Current Region (H. G. Moser, ed.), p. 1369-1403. CalCOFI Atlas 33.

Cooper, J., and F. Chapleau. 1996. The monophyletic status of Pleuronectes (Pleuronectidae: Pleuronectiformes). In Abstracts of the 1996 Annual Meeting of the American Society of Ichthyologists and Herpetologists, New Orleans, LA, p. 114.

1998. Monophyly and intrarelationships of the family Pleuronectidae (Pleuronectiformes), with a revised classification. Fish. Bull. 96:686-726.

Cope, E. D. 1873. A contribution to the ichthyology of Alaska. Proc. Am. Phil. Soc. 13:24-32.

Environmental Systems Research Institute. 1996. Arcview GIS, ver. 3.0. Environmental Systems Research Institute, Redlands, CA, 614 p.

Evermann, B. W., and E. L. Goldsborough. 1907. The fishes of Alaska. Bull. Bur. Fish. 26:219-360.

Fadeev, N. S. 1965. Comparative outline of the biology of flatfishes in the southeastern part of the Bering Sea and condition of their resources. Soviet Fisheries Investigations in the northeast Pacific, Part 4. Pacific Scientific Research Institute of Marine Fisheries and Oceanography (TINRO). Izvestiya 58:112-129. [1968 transl. by Israel Prog. Sci. Transl.]

Forrester, C. R. 1969. Life history information on some groundfish species. Fish. Res. Board Can. Tech. Rep. 105, 17 p.

Forrester, C. R., and J. A. Thompson. 1969. Population studies on the rock sole (Lepidopsetta bilineata (Ayres)) of northern Hecate Strait, B.C. Fish. Res. Board Can., Tech. Rep. 108, 104 p.

Garrison, K. J., and B. S. Miller. 1982. Review of the early life history of Puget Sound fishes. Report FRI-UW-8216 of the Fish. Res. Inst., Univ. Washington, Seattle, WA, 729 p.

Gill, T. N. 1862. Note on some genera of fishes of western North America. Proc. Acad. Nat. Sci. Phila. 14:329-332.

1864. Synopsis of the pleuronectoids of California and northwestern America. Proc. Acad. Nat. Sci. Phila. 16:194-198.

Girard, C. F. 1854. Descriptions of new fishes, collected by Dr. A. L. Heermann, naturalist attached to the survey of the Pacific railroad route, under Lieut. R. S. Williamson, U.S.A. Proc. Acad. Nat. Sci. Phila. 7:129-140.

1856. Contributions to the ichthyology of the western coast of the United States, from specimens in the collection of the Smithsonian Institution. Proc. Acad. Nat. Sci. Phila. 8:131-137.

Goode, G. B. 1884. The fisheries and fishery industries of the United States. Section I. Natural history of useful aquatic animals with an atlas of two hundred and seventy-seven plates. Plates. Government Printing Office, Washington, D.C., 215 p.

Gunther, A. 1862. Catalogue of the fishes in the British Museum. Catalogue of the Acanthopterygii Pharyngognathi and Anacanthini in the collection of the British Museum, vol. 4, 534 p.

Hart, J. L. 1973. Pacific fishes of Canada. Bull. Fish. Res. Board Can. 180, 740 p.

Hickman, C. P., Jr. 1959. The larval development of the sand sole (Psettichthys melanostictus). Wash. Dep. Fish. Res. Pap. 2:38-47.

Hubbs, C. L. 1915. Flounders and soles from Japan collected by the United States Bureau of Fisheries steamer “Albatross” in 1906. Proc. U.S. Natl. Mus. 48:449-496.

Hubbs, C. L., and K. F. Lagler. 1958. Fishes of the Great Lakes region. Cranbrook Inst. Sci., Ann Arbor, MI, 213 p.

Ivankov, V. N. 1996. Review of the book by G. U. Lindberg and V. V. Fedorov, The fishes of the Sea of Japan and adjacent parts of the Sea of Okhotsk and the Yellow Sea. Part 6. Teleostomi, Osteichthyes, Actinopterygii. XXXI. Pleuronectiformes–Family Psettodidae–CCICFamily Cynoglossidae. J. Ichthyol. 36(7):549-550. [Transl. by P. A. Aleinikov.]

Jordan, D. S., and B. W. Evermann. 1898. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part III. Bull. U.S. Natl. Mus. 47:2183-3136.

1900. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part IV. Bull. U.S. Natl. Mus. 47:3137-3313.

Jordan, D. S., and C. H. Gilbert. 1881. List of the fishes of the Pacific Coast of the United States, with a table showing the distribution of the species. Proc. U.S. Natl. Mus. 3:452-458.

Jordan, D. S., and D. K. Goss. 1889. A review of the flounders and soles (Pleuronectidae) of America and Europe. Rep. U.S. Fish Comm. 14:225-342.

Jordan, D. S., and C. L. Hubbs. 1925. Record of fishes obtained by David Starr Jordan ia Japan, 1922. Mem. Ann. Carnegie Mus. 10(2):93-346.

Jordan, D. S., and E. C. Starks. 1906. A review of the flounders and soles of Japan. Proc. U.S. Natl. Mus. 31(1484):161-246.

Kabata, Z. 1988. Copepoda and Branchiura. In Guide to the parasites of fishes of Canada, part II– Crustacea (L. Margolis and Z. Kabata, eds.), p. 3-127. Can. Spec. Publ. Fish. Aquat. Sci. 101.

Kendall, A. W., Jr., E. H. Ahlstrom, and H. G. Moser. 1984. Early life history stages of fishes and their characters. In Ontogeny and systematics of fishes (H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall Jr., and S. L. Richardson, eds.), p. 11-22. Am. Soc. Ichthy. Herpet. Spec. Publ. 1.

Kim, I. S., and C. H. Youn. 1994. Taxonomic revision of the flounders (Pisces: Pleuronectiformes) from Korea. Korean J. Ichthy. 6(2):99-131.

Kravitz, M., W. G. Pearcy, and M. P. Guin. 1976. Food of five species of co-occurring flatfishes on Oregon’s continental shelf. Fish. Bull. 74:984-990.

Lang, G. M., P. A. Livingston, and B. S. Miller. 1995. Food habits of three congeneric flatfishes: yellowfin sole (Pleuronectes asper), rock sole (P. bilineatus), and Alaska plaice (P. quadrituberculatus) in the eastern Bering Sea. Proc. InterNatl. Symp. N. Pac. Flatfish, Alaska Sea Grant Prog. Report AK-SG-95-04, p. 225-245.

Levings, C. D. 1967. A comparison of growth rates of the rock sole (Lepidopsetta bilineata) Ayres, in northeast Pacific waters. Fish. Res. Board Can., Tech. Rep. 36, 43 p.

Leviton, A. E., and M. L. Aldrich. 1997. Theodore Henry Hittell’s “The California Academy of Sciences, a narrative history: 1853-1906.” Calif. Acad. Sci., San Francisco, CA, 623 p.

Leviton, A. E., and R. H. Gibbs Jr. 1988. Standards in herpetology and ichthyology. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Suppl. 1: additions and corrections. Copeia 1988:280-282.

Leviton, A. E., R. H. Gibbs Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832.

Lindberg, G. U., and V. V. Fedorov. 1993. The fishes of the Sea of Japan and adjacent parts of the Sea of Okhotsk and the Yellow Sea. Part 6. Teleostomi, Osteichthyes, Actinopterygii. XXXI. Pleuronectiformes–Family Pleuronectidae. Zool. Inst. Rossiiskoi Akad. Nauk, 271 p.

Lockington, W. N. 1879a. The flounders of our markets, No. 2. Mining and Scientific Press 39(4):70.

1879b. Review of the Pleuronectidae of San Francisco. Proc. U.S. Natl. Mus. 2:69-108.

1880a. Note on a new flat-fish (Lepidopsetta isolepis) found in the markets of San Francisco. Proc. U.S. Natl. Mus. 3:325.

1880b. Notes on new and rare fishes of the Pacific Coast. Am. Nat. 14:595-600.

1883. Isopsetta isolepis. In 1883 synopsis of the fishes of North America (D. S. Jordan and C. H. Gilbert, eds.), p. 1-1018. Bull. U. S. Natl. Mus. 16.

Manugistics. 1997. Statgraphics plus 2.1. Manugistics, Inc., Rockville, MD, 882 p.

Masutomi, K., and T. Hamada. 1966. Fossils in colour. Hoikusha Publishing Co., Osaka, 268 p. [Not seen; cited in Sakamoto and Uyeno (1988).]

Matarese, A. C., A. W. Kendall Jr., D. M. Blood, and B. M. Vinter. 1989. Laboratory guide to early life history stages of northeast Pacific fishes. U.S. Dep. Commer., NOAA Tech. Rep., NMFS 80, 652 p.

Minami, T. 1995. Spawning season of pleuronectid flatfishes in the coastal waters of Hokkaido, Japan [Review]. Bull. Hokkaido Natl. Fish. Res. Inst. 59:69-80.

Minami, T., and I. Nakamura. 1978. Seasonal occurrence of pelagic stages of flatfishes in the coastal waters of western Wakasa Bay. Mem. Coll. Agriculture, Kyoto Univ. 112:29-47.

Moiseev, P. A. 1953. [Cod and flounders of Far-Eastern Seas.] Izv. Tikhook. N.-I. Inst. Rybn.-Khoz. I Okeanogr. 40:1-287. Fish. Res. Board Can. Transl. 119.]

Moser, H. G., W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall Jr., and S. L. Richardson, eds. 1984. Ontogeny and systematics of fishes. Am. Soc. Ichthy. Herpet. Spec. Pub]. 1,760 p.

Moser, H. G., R. L. Charter, P. E. Smith, D. A. Ambrose, S. R. Charter, C. A. Meyer, E. M. Sandknop, and W. Watson. 1993. Distributional atlas offish eggs and larvae in the California Current region: taxa with 1000 or more total larvae, 1951 through 1984. CalCOFI Atlas 31, 233 p.

Mulligan, H. L., A. W. Kendall Jr., and A. C. Matarese. 1995. The significance of morphological variation in adults and larvae of the rock sole, Pleuronectes bilineatus, from the Bering Sea and northeastern Pacific Ocean. Proc. Int. Symp. N. Pac. Flatfish, Alaska Sea Grant Coll. Prog. Report AK-SG-95-04, p. 133-150.

NMFS (National Marine Fisheries Service). 1999. Our living oceans: report on the status of U.S. living marine resources, 1999. U.S. Dep. Commer., NOAA Tech. Memo. NMFS-F/SPO-41, 301 p.

Nelson, J. S. 1994. Fishes of the world, 3rd ed. Wiley & Sons, New York, NY, 600 p.

Nikiforov, S. N., S. N. Safronov, and N. S. Fadeev. 1983. Distinguishing characters of rock sole, Lepidopsetta bilineata, and mochigar sole, L. mochigarei (Pleuronectidae). J. Ichthyol. 23:36-44.

Norman, J. R. 1934. A systematic monograph of the flatfishes (Heterosomata). Brit. Mus. Nat. Hist. 1:1-459.

Okada, Y. 1955. Fishes of Japan, illustrations and descriptions of fishes of Japan. Maruzen Co., Tokyo, 434 p.

Okiyama, M., ed. 1988. An atlas of the early stages of fishes in Japan. Tokai Univ. Press, Tokyo, 1154 p. [In Japanese.]

Okiyama, M., and K. Takahashi. 1976. Larval stages of right eye founders (subfamily Pleuronectinae) occurring in Japan Sea. Bull. Japan. Sea. Reg. Fish. Res. Lab. 27:11-34.

Penttila, D. E. 1995. The WDFW’s Puget Sound intertidal baitfish spawning beach survey project. In Puget Sound Research 95 Proceedings, vol. 1, Jan. 12-14, 1995, p. 235-241.

Pertseva-Ostroumova, T. A. 1961. The reproduction and development of far eastern flounders. Tr. Inst. Okeanol. Akad. Nauk SSR. [In Russian; English transl., Fish. Res. Board Can., Transl. Serv. 856, 1967.]

Poss, S. G., and B. B. Collette. 1995. Second survey of fish collections in the United States and Canada. Copeia 1995:48-70.

Rass, T. S. 1996. On taxonomy of Pleuronectini (Pleuronectidae). J. Ichthyol. 36:546-548.

Sakamoto, K. 1984a. Interrelationships of the family Pleuronectidae (Pisces: Pleuronectiformes). Mem. Fac. Fish., Hokkaido Univ. 31(1/2):95-215.

1984b. Pleuronectidae. In The fishes of the Japanese Archipelago (H. D. Masuda, K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino, eds.), p. 351-354. Tokai Univ. Press, Tokyo, Japan.

Sakamoto, K., and T. Uyeno. 1988. A new righteye flounder from the late Pleistocene Togane Formation, Chiba Prefecture, Japan. Bull. Nat. Sci. Mus., Tokyo, ser. C, 14(3):135-142.

Schmidt, P. Y. 1950. Fishes of the Sea of Okhotsk. Academy of Sciences of the USSR, Transactions of the Pacific Committee, vol. 6, Moscow-Leningrad, 392 p. [In Russian, 1965 transl. by Israel Prog. Sci. Trans. Available from the National Technical Information Service, Springfield, VA.]

Shubnikov, D. A., and L. A. Lisovenko. 1964. Data on the biology of rock sole of the southeastern Bering Sea. Soviet Fisheries Investigations in the northeast Pacific, Part 2. Pacific Scientific Research Institute of Marine Fisheries and Oceanography (TINRO). Izvestiya 49:220-226. [1968 transl, by Israel Prog. Sci. Transl.]

Shvetsov, F. C. 1979. Reproduction of the flounder Lepidopsetta bilineata bilineata, off the Okhotsk Sea coast near Paramoshiro and Shumushu Islands. J. Ichthyol. 19(5):61-62.

Skalkin, V. A. 1959. [Diet and food relations of flatfish on the II’inskii Shoal.] Pacific Scientific Research Institute of Marine Fisheries and Oceanography (TINRO). Izvestiya 47. [In Russian; cited by Skalkin, 1963.]

1963. Diet of flatfishes in the southeastern Bering Sea. Soviet Fisheries Investigations in the Northeast Pacific, Part 1. Pacific Scientific Research Institute of Marine Fisheries and Oceanography (TINRO). Izvestiya 50:235-250. [1968 transl, by Israel Prog. Sci. Transl.]

Smith, R. T. 1936. Report on the Puget Sound otter trawl investigations. Wash. Dept. Fish. Biol. Rep. 36B:1-61.

Snyder, J. O. 1911. Descriptions of new genera and species of fishes from Japan and Riu Kiu Islands. Proc. U.S. Natl. Mus. 40:525-549.

1912. Japanese shore fishes collected by the United States Bureau of Fisheries Steamer “Albatross” expedition of 1906. Proc. U.S. Natl. Mus. 42:437-441.

Sokal, R. R., and F. J. Rohlf. 1995. Biometry, 3rd ed. W.H. Freeman and Co., New York, NY, 859 p.

Springer, V., and M. E. Anderson. 1998. Catalog of type specimens of recent fishes in the National Museum of Natural History, 8: Suborder Zoarcoidei (Anarhichadidae, Bathymasteridae, Pholidae, Ptilichthyidae, Scytalinidae, Stichaeidae, Zoarcidae). Smiths. Contr. Zool. 589, 27 p.

SPSS, Inc. 1996. SYSTAT for Windows, version 6.0. SPSS, Inc., Chicago, IL, 751 p.

Statistical Sciences. 1993. S-Plus for Windows, version 3.3. Statistical Sciences, Seattle, WA, 819 p.

Stauffer, J. R., Jr., and E. Hert. 1992. Pseudotropheus callainos, a new species of mbuna (Cichlidae), with analyses of changes associated with two intra-lacustrine transplantations in Lake Malawi, Africa. Ichthyol. Explor. Freshwaters 3:253-264.

Taranets, A. J. 1937. Handbook for identification of fishes of the Soviet Far East and adjacent waters. Bull. Pac. Sci. Fish. Inst. Vladivostok. 11:1-200+map.

Townsend, L. D. 1936. Variations in the meristic characters of flounders from the northeastern Pacific. Rep. Intern. Fish. Comm. 11:1-24.

1937. Geographical variation and correlation in Pacific flounders. Copeia 1937:92-103.

Wilimovsky, N. J., A. Peden, and J. Peppar. 1967. Systematics of six demersal fishes of the North Pacific Ocean. Fish. Res. Board Can. Tech. Rep. 34, 95 p.

Yusa, T. 1957. Eggs and larvae of flatfishes in the coastal waters of Hokkaido. II. Early development of the flatfish (Lepidopsetta mochigarei Snyder). Bull. Hokkaido Reg. Fish. Res. Lab. 15(2):15-21.

1958. Eggs and larvae of flatfishes in the coastal waters of Hokkaido. II. Post-embryonic development and larvae of the flatfish Lepidopsetta mochigarei Snyder. Bull. Hokkaido Reg. Fish. Res. Lab. 18:1-10.

Appendix Table 1 Summary of Alaska Fisheries Science Center cruises with positive tows for larvae of Lepidopsetta (1971-72, 1977-94). Cruises in boldface were not used in calculations of mean density. BS = Bering Sea; GA = Gulf of Alaska; WA = Washington. Bongo = bongo net of 60 cm diameter, 0.505-mm mesh (larvae were occasionally collected using alternative frame sizes, including 20-cm net diameter and 0.333-mm mesh, but these collections were not used for abundance calculations); Meth = Methot trawl net of 5-m2 frame (2 x 3 mm oval mesh); Tuck = Tucker trawl net of 1-m frame and 0.505-mm mesh.

Year Cruises Region Month Gear(1)

1971 4DE71 BS Jul-Aug bongo

1972 2KE72 GA Apr-May bongo

1977 MF77B-5,6 BS Apr-May bongo

9SEI77 GA Jul bongo

1978 4DI78 GA Mar-Apr bongo, Tuck

2MF78 GA Jun-Jul bongo, Tuck

3MF78 GA Sep bongo

4MF78 GA Sep-Oct bongo

5MF78 GA Oct-Nov bongo

1979 1PO79 GA Sep bongo

3MF79 BS Jun-Jul bongo, Tuck

5TK79 GA May bongo

1980 1MF80 BS Jan-Feb bongo

1981 1PO81 WA May-Jun bongo

1SH81 GA Mar bongo

2MF81 GA Mar-Apr bongo

2SH81 GA Apr bongo

3MF81 GA Apr-May bongo

3SH81 GA May bongo

4MF81 GA May bongo

1982 1DA82 GA Apr bongo

2DA82 GA May bongo

1983 1CH83 GA May bongo

1EQ83 WA Apr-May bongo

1984 1SH84 GA Apr-May bongo

1985 1PO85 GA Mar-Apr bongo

2MF85 GA May bongo

2PO85 GA May-Jun bongo

1986 1GI86 GA Apr bongo

1MF86 GA Apr bongo

2MF86 GA May bongo

1987 1BB87 GA Mar-Apr bongo

2MF87 GA Apr bongo

3MF87 GA May bongo

1988 1DN88 BS and GA Mar-May bongo

1MF88 GA Apr bongo, Tuck

2MF88 GA Apr-May bongo, Meth, Tuck

3MF88 GA May bongo

4MF88 GA May-Jun bongo, Meth, Tuck

1989 1MF89 GA Apr bongo, Tuck

2MF89 GA Apr-May bongo, Tuck

3MF89 GA May bongo, Meth, Tuck

4MF89 GA May-Jun Tuck

1990 1MF90 GA Apr bongo

2MF90 GA May bongo, Tuck

3MF90 GA May bongo

4MF90 GA May-Jun bongo

1991 1MF91 GA Apr bongo, Tuck

2MF91 GA Apr bongo, Tuck

3MF91 GA May bongo

4MF91 GA May bongo

5MF91 WGA July Meth

1992 1MF92 GA Apr bongo, Tuck

1MM92 BS and GA Jul Meth

3MF92 GA May bongo, Tuck

4MF92 GA May bongo, Meth, Tuck

1993 2MF93 GA Apr bongo, Meth, Tuck

3MF93 BS Apr bongo, Tuck

4MF93 GA May bongo, Meth, Tuck

5MF93 GA May-Jun bongo, Meth

6MF93 GA Sep Tuck

1994 3MF94 GA Mar-Apr bongo, Tuck

4MF94 BS Apr bongo, Tuck

Total occurrences

L. polyxystra

Cruises Total stations n. sp. L. bilineata

4DE71 10 8(2) 0

2KE72 67 27 0

MF77B-5,6 75 35 0

9SEI77 99 0 1(2)

4DI78 225 80 0

2MF78 267 41 66

3MF78 26 1 1

4MF78 66 2 0

5MF78 19 0 1

1PO79 18 0 1

3MF79 271 35 0

5TK79 58 15 3

1MF80 8 2(2) 0

1PO81 131 0 1

1SH81 131 4 0

2MF81 89 1 0

2SH81 60 31 2

3MF81 79 37 2

3SH81 57 37 10

4MF81 80 52 8

1DA82 83 20 0

2DA82 62 29 1

1CH83 70 19 5

1EQ83 124 0 1

1SH84 157 22 1

1PO85 154 37 0

2MF85 62 4 0

2PO85 189 52 17

1GI86 149 53 0

1MF86 80 4 0

2MF86 107 53 0

1BB87 117 12 1

2MF87 142 9 0

3MF87 60 14 8

1DN88 203 17 0

1MF88 196 51 0

2MF88 84 41 1

3MF88 13 2 0

4MF88 209 69 75

1MF89 140 21 0

2MF89 107 55 1

3MF89 221 127 2

4MF89 95 87 9

1MF90 107 17 0

2MF90 92 52 2

3MF90 17 9 0

4MF90 130 83 21

1MF91 98 5 0

2MF91 156 24 0

3MF91 119 28 0

4MF91 98 19 8

5MF91 62 40 12

1MF92 103 7 0

1MM92 11 3 2

3MF92 188 59 7

4MF92 154 46 31

2MF93 135 12 0

3MF93 135 21 0

4MF93 168 44 7

5MF93 119 25 22

6MF93 24 0 2

3MF94 49 2 0

4MF94 144 37 0

(1) Although larvae of Lepidopsetta were collected in neuston and Mocness tows (see “Material examined” section), these collections were not used for abundance calculations and are not listed here.

(2) Total occurrences for these cruises represent the minimum number collected. Complete records for these cruises are unavailable at this time.

James W. Orr

Ann C. Matarese

Resource Assessment and Conservation Engineering Division

Alaska Fisheries Science Center

National Marine Fisheries Service, NOAA

7600 Sand Point Way NE

Seattle, Washington 98115-0070

E-mail address (for J. W. Orr): James.Orr@noaa.gov

Manuscript accepted 22 February 2000.

COPYRIGHT 2000 National Marine Fisheries Service

COPYRIGHT 2004 Gale Group