DESCRIPTION OF TilE LARVAL STAGES OF FIVE NORTHERN CALIFORNIA SPECIES OF ROCKFISHES (FAMILY SCORPAENIDAE) FROM REARING S1UDIES

A Thesis presented to the Faculty of California State University, Stanislaus and Moss Landing Marine Laboratories

In Partial Fulfillment Of the Requirements for the Degree Master of Science in Marine Science

by Guillermo Moreno August, 1990 ABSTRACT

There are about 72 species of (Family Scorpaenidae) along the eastern Pacific coast of North America, some of which are heavily exploited by both commercial and sport fisheries. Larval identifications are needed for management-related studies of these fishes. However, due to the large number of species and recent evolutionary divergence in this genus, the identification of early life stages has progressed slowly. In this study, four species of rockfish (Sebastes mystinus, S. camatus, S. atrovirens, and S. rastrelliger) were reared and described, and larvae of another (S. melanops) reared elsewhere were described. The larvae were fed a mix of , brine shrimp (Artemia) nauplii, and daily-caught plankton. Of the descriptions of eastern Pacific Sebastes larvae to date, only nine have been based on reared larvae, due to the difficulty in rearing them past the yolk absorption stage. Two general pigmentation patterns were discerned: (1) a short row of ventral midline melanophores on the trunk, and no or very little posterio-dorsal pigmentation (S. mystinus and S. melanops); and (2) complete ventral midline pigmentation on the trunk, and anterior and posterio-dorsal melanophores (S. carnatus, S. atrovirens, and S. rastrelliger). With the exception of very early stages of S. carnatus and S. atrovirens, these five species can be distinguished from each other based

11 on pigmentation characteristics. The morphometric proportions did not demonstrate major differences among species. Although difficult, identification of the larval stages of Sebastes spp. is possible for most of the species described to date. Culture and descriptive techniques need to be applied to other species of Sebastes to help reduce the number of factors that confuse the of this complex group.

iii ACKNOWLEDGEMENTS

First, I would like to thank my parents Maria Elena and Ricardo Moreno for their moral support and encouragement through these years. I owe many thanks to my committee members, Drs. G. M. Cailliet, V. Loeb, and P. Roe for their support and interest in my work, and for critically reviewing this manuscript The final draft of this thesis would not have been possible without their comments and ideas. Many people helped in this project, however, I cannot mention all of them. Suffice it to say that I am indebted to all of them. Many thanks are due to L. McMasters who drew the larvae and helped many times with drafting and photographic work. I would like to thank C. Miller for providing the S. melanops larvae. Food rearing techniques were improved with the help of R. Orhun at Hubbs/Sea World Research Institute. This research would not have been possible without the help and collaboration of the Monterey Bay Aquarium. I am grateful to the Husbandry Department at the MBA for its continuous interest, and logistical support. Many thanks are due to the faculty, staff, and students of Moss Landing Marine Laboratories. Their help and encouragement have been extraordinary. This work is the result of research sponsored in part by NOAA,

lV National Sea Grant College Program, Department of Commerce, under grant number NA85AA-D-56140, project number R/F-115, through the California Sea Grant College Program, and in part by the California State Resources Agency. The U.S. Government is authorized to reproduce and distribute for governmental purposes. This grant was awarded to Drs. G. M. Cailliet, and V. Loeb. Additional support was given to the author by the David and Lucile Packard Foundation.

v TABLE OF CONTENTS

Page

ABS1RACT 11

ACKNOWLEDGEMENTS IV

THESIS APPROVAL VI

LIST ofTABLES IX

LIST of FIGURES X INTRODUCTION 1 METHODS 5 Collection 5 Rearing 5 Descriptions 7 RESULTS 10 Descriptiom: 10 Sebastes mystinus 11 Sebastes carnatus 12 13 14 Sebastes melanops 15 Species Comparisons 17 a. Morphometries 17 b. Pigmentation 18 DISCUSSION 20 LITERATURE CITED 27

Vll TABLES 34 FIGURES 44

viii LIST OF TABLES Page

1. Proportions of Sebastes mystinus larvae with melanophores present at loci 1-33. 34

2. Proportions of Sebastes camatus larvae with melanophores present at loci 1-33. 35

3. Proportions of Sebastes atrovirens larvae with melanophores present at loci 1-33. 36

4. Proportions of Sebastes rastrelliger larvae with melanophores present at loci 1-33. 37

5. Proportions of Sebastes melanops larvae with melanophores present at loci 1-33. 38

6. Morphometric proportions of Sebastes mystinus larvae. 39

7. Morphometric proportions of Sebastes camatus larvae. 40

8. Morphometric proportions of Sebastes atrovirens larvae. 41

9. Morphometric proportions of Sebastes rastrelliger larvae. 42 10. Morphometric proportions of Sebastes melanops larvae. 43

IX LIST OF FIGURES Page 1. Geographic distributions of adult Sebastes mystinus, S. carnatus, S. atrovirens, S. rastrelliger, and S. melanops along the west coast of North America. 44

2. Capture site of gravid Sebastes mystinus and S. carnatus females. Rearing of all species butS. melanops occurred at the Monterey Bay Aquarium. 46

3. Diagram of Sebastes larva, with axes used for morphometric measurements. 48

4. Diagram of Sebastes larva, with map of melanophore loci locations. 50

5. Starved Sebastes mystinus larva. 52

6. Growth curves of Sebastes mystinus, S. camatus, S. atrovirens, S. rastrelliger, and S. melanops larvae. 54

7. Growth series of Sebastes mystinus larvae. 56

8. Melanophore densities (mean number per larva) for loci 4 and 6 for all species. 58

9. Melanophore densities (mean number per larva) for loci 18 and 19 for all species 60

10. Growth series of Sebastes carnatus larvae. 62

11. Growth series of Sebastes atrovirens larvae. 64

12. Growth series of Sebastes rastrelliger larvae. 66

13. Growth series of Sebastes melanops larvae. 68

X 1

INTRODUCTION

The genus Sebastes (Family Scorpaenidae) is an abundant and diverse group of fishes along the Pacific coast of North America (Love and Westphal, 1981). It includes about 72 species and 11 subgenera (Kendall, in press), and is a very important part of the commercial and sport fishery catches off California, Oregon, Washington, Canada, and Alaska (Moser et. al., 1977; Gunderson and Lenarz, 1980; Lenarz, 1987). Rockfishes comprise one third of the recreational fish catch in California , with S. mystinus, S. melanops, and S. flavidus comprising up to 30% of this proportion. In addition, S. mystinus and S. melanops are also important components of the commercial rockfish catch (Lenarz, 1987). Due to increasing fishing pressures and to the lack of information on spawning stock size and recruitment relationships, additional information on life history characteristics is needed to foster comprehensive management of rockfish fisheries. An example of the sensitivity to overfishing of species in this group is the (Sebastes alutus) in the northeast Pacific. Dramatic drops in the production and catch per unit effort due to increased fishing pressure during the 1960's, and late 1970's through the early 1980's were seen (Haldorson, 1987) . Although relatively fecund, the Pacific ocean perch is characterized by having a relatively advanced age at maturity (9-11 years for females, and 6-7 years for males; Gunderson, 1977), a trait common within this genus (Wyllie-Echeverria, 1985). lchthyoplankton studies provide information for the detection and appraisal of fishery resources, and studies of biology, systematics, and population dynamics of fishes (Hempel, 1973). The early life stages of fishes are usually restricted to the upper several hundred meters of the water column, 2 and for most marine fishes are either passive eggs or weakly swimming larvae, making it possible to sample a variety of species over a broad range with plankton nets (Smith and Richardson, 1977). Species identification of early life history stages is important for ichthyoplankton studies. In the genus Sebastes, many characters are shared and many of the species are very closely related, making identification and evaluation on the importance of characters very difficult (Moser et al., 1977; Moser and Ahlstrom, 1978; Barsukov, 1981; Kendall and Lenarz, 1987). Larval rearings provide the essential identification information and also allow for an evaluation of the range of variation of characters demonstrated by larvae from various ages, broods, and species. This allows determination of the relative importance of various diagnostic characters, essential for studies of the genus Sebastes. Rockfishes are viviparous, giving birth from a few hundred thousand to over two million larvae (e.g., S. paucispinis; Moser, 1967) during an annual spawning season. The newly extruded larvae are born with little yollc, but with well developed eyes and mouths, allowing immediate feeding on live food (Boehlert and Yoklavich, 1984). However, when compared to larvae of similar developmental stages of other genera, these larvae are not as capable of capturing large-size prey (Bainbridge and McKay, 1968). Partial descriptions are available for the larvae of 51 eastern Pacific larval rockfish species, leaving a large portion of the genus Sebastes, including important commercial and recreational species, still undescribed. Because of the relatively recent speciation and extensive radiation of the genus (Moser et al., 1977; Moser and Ahlstrom, 1978; Barsukov, 1981; Kendall and Lenarz, 1987) identification of larval, juvenile and adult stages can be difficult. Early developmental stages traditionally have been described by 3 using preserved field collected specimens and working backwards from newly transformed, identifiable juveniles to larval stages (Morris, 1956; Moser, 1967, 1972; Waldron, 1968; Westrheim, 1975; Moser, eta!. 1977; Moser and Ahlstrom, 1978; Richardson and Laroche, 1979; Laroche and Richardson, 1981; Moser, eta!. 1985). However, a more reliable and accurate technique involves rearing larvae from known parental stocks through caudal fm formation and later stages if possible (Moser and Butler, 1981; Stahl-Johnson, 1984, 1985; Moser and Butler, 1987; Wold, 1990). Rearing oflarval Sebastes has met with varying success. Workers in the northwestern Pacific have raised larvae of eight species, three of these to the juvenile stage (S. pachycephalus, S. oblongus, and S. schlegeli: Fujita, 1957, 1958; Siokawa and Tsukahara, 1961; Talcai and Fukunaga, 1971; Hoshiai, 1977; Kusakari eta!., 1977). However, only recently have eastern Pacific Sebastes larvae been raised past yolk absorption and in some cases up to caudal fin formation (Moser and Butler, 1981, 1987; Stahl-Johnson, 1985; Wold, 1990). Factors responsible for difficulties in rearing the northeastern Pacific species include their comparatively small size at birth (<4.5 mm) and cool ambient water temperatures that retard growth rates relative to western Pacific species (Kendall and Lenarz, 1987). The purpose of this study was: (1) to rear the larvae of various rockfish species which are widely distributed along the west coast of North America (Eschmeyer et. a!., 1983) (Figure 1) and are associated with beds and/or nearshore rocky reef areas; and {2) to describe their sequential stages of early development. The species studied were the (Sebastes mystinus), (S. carnatus), kelp rockfish (S. atrovirens), and grass rockfish (S. rastrelliger). In addition, the newborn and three day old larvae of the (S. melanops) were described from 4 material provided by Telonicher Marine Laboratory (Humboldt State University, Arcata, CA). 5

:METHODS

Collection

Gravid females of Sebastes mystinus (n = 4) and S. carnatus (n = 3) were collected at Stillwater Cove, Carmel Bay (Figure 2) between December­ May 1988 and1989. Advanced pregnancy in all females was evident from the distended abdomens and vent areas, thereby permitting selective capture. Rockfishes were captured by SCUBA divers using hook and line or gillnets, transported to the Monterey Bay Aquarium (MBA), and placed in rearing tanks until after parturition. Sebastes atrovirens (n = 2) and S. rastrelliger (n = 1) larvae were obtained from gravid females residing in the Kelp Tank at the MBA. Food (squid, krill, and anchovies) was offered to all females during this period, but no feeding was observed.

Rearing

The rearing tanks were 400 L, with black walls and white conical bottoms, and had a continuous flow (3-4 L/min) offiltered (1 11m) seawater. The flowthrough system was similar to that of Hughes eta!. (1974), differing only in the direction of the flow, which in this study was from top to bottom. At pa1turition, or soon after, the females were removed from the rearing tanks, and larval densities were reduced to <10 larvae/Las recommended by Stahl-Johnson (1985), by siphoning the excess larvae until desired densities were reached. Densities were evaluated visually. The larvae of S. mystinus, S. atrovirens, S. carnatus, and S. rastrelliger were provided with a variety of food organisms. Larvae up to one week old were fed 3-4 times per day with 6

the marine plicatilis and wild plankton. The plankton was collected daily in Elkhorn Slough, Monterey County, CA, with a 30 Jll11 net, and filtered to 120 J..lffi. Possible prey items observed in the plankton included rotifers, copepods, nauplii, tintinnids, trochophore larvae, and various algal species, mainly diatoms. After the first week, brine shrimp (Artemia spp.) nauplii (San Francisco Bay Brand, Inc., Newark, CA ) were added to the diet. The wild plankton was then filtered to 220 J..lffi to allow larger prey items in the tanks, but to avoid introduction of possible predators and other fish larvae. The reared rotifers were fed cultured algae (Tetraselmus spp., Phaeodactylum spp., lsochrysis spp., and Duniolella spp.; Guillard, 1975) and were enriched daily with a mixture of fish oil (herring) and egg yolk (1:1) (R. Orhun, Hubbs Marine Research Center, San Diego, CA, pers. comm.). The Artemia nauplii were enriched with "Selco" (Artemia Systems N.Y., Belgium), and the wild plankton was treated with a fungicide-bactericide (1- 10 ppm; "Prefuran", Argent Labs., Redmond, W A) to reduce the possibility of bacterial contamination of the larval rearing tanks. The latter measure was taken after large pink bacterial blooms (Pseudomonas spp.) were observed in the rearing tanks. During the first rearing season, water quality samples were taken weekly and analyzed for nitrate, nitrite, phosphate, ammonia, and oxygen levels. Due to consistently high quality of the water system, these tests were later discontinued. Continuous lighting was introduced to allow around-the-clock feeding, to better the odds of survival and growth. Sebastes melanops larvae, extruded from a field-caught female, were reared by Chris Miller at the Telonicher Marine Laboratory. Due to the lack of growth and starvation in these larvae, only one to three day old specimens were described. 7

Descriptions

Larval fish were sampled every four to five days for descriptive work, photographs and preservation. When possible, the larvae were chilled to slow activity for photography. However, most photographs used for subsequent illustrations were derived from previously preserved larvae due to the delicate nature and rapid decomposition of the larvae prior to preservation. Larvae initially were fixed in 5% buffered (sodium borate) formalin, and transferred after 30 days to a dehydrating series of isopropanol (25-50%) and ethanol (70% ), allowing a week between each alcohol change. To describe the Sebastes larvae it was necessary to choose a character, either age or size, to allow an evaluation of variance, and a determination of useful specific identifying characters. In this study, size was chosen as the main character to separate larvae and allow comparisons. Morphometric and pigmentation analyses were conducted on dehydrated larvae for 0.5 mm size classes within the available size range (e.g. 3.0-7.9 mm) for each species. The morphometric measurements included: standard length (SL); snout-anus length (SA); head length (HL); snout length (SnL); eye diameter (ED); and body depth at pectoral fin base (BD) (Figure 3; Moser et al., 1977; Moser and Ahlstrom, 1978). Body proportions (SA/SL; HL/SL; Snl)HL; ED/HL; BD/SL) were calculated and means, standard deviations, and ranges were obtained for each size group of each species. Pectoral fin length measurements were made only on the older (8-31 days) S. mystinus larvae, due to the difficulty in obtaining reliable measurements for the other stages of all five species. The pectoral fin length measurements for younger S. mystinus and all sizes of the other four species 8 were of dubious precision due to the lack of pigmentation on the pectoral fin outer margins, a character that delineated the fins and permitted reliable measurements. In this study, a modification of the pigment presence/absence scheme of Kendall and Lenarz (1987), which utilized 26 loci, and marked the presence/absence of melanophores without indicating what proportion of larvae had pigmentation on those loci, was used. Pigmentation analysis of 33 loci (Figure 4) assessed the presence and number of melanophores for loci 4, 6, 18, and 19, and the presence (given as proportions) of melanophores for the rest of the loci. Loci 4, 6, 18, and 19 were treated differently because they allowed melanophore counting with confidence. The other loci had tightly packed melanophores that made counts impossible in most cases. Due to large observed variability in melanophore development, proportions of larvae with pigment presence at each locus and size class were noted in this study. Common pigmentation sites in Sebastes larvae include the dorsal and lateral surfaces of the gut (loci ll, 12, 15, and 17), dorsal midline of the tail (loci 21-23), pectoral fins (loci 7 -9), nape (locus 4 ), dorsal aspect of the head (locus 6), upper and lower jaws (loci 1-3, and 29), and caudal fin (loci 20, 25 and 26; Kendall and Lenarz, 1987). Pigmentation on the forehead (locus 4), nape (locus 6), and dorsal midline (loci 18 and 19) served to distinguish between the species treated here. Means, standard deviations, and ranges of melanophore numbers at these loci were calculated for each 0.5 mm size class. With the exception of S. mystinus and S. melanops, the number of melanophores in loci 22 and 23 were not counted due to their generally dense arrangement.

Statistics (ANOVAs) were utilized in an attempt to compare morphometric measurements and pigmentation patterns among species. 9

However, due to the small number of females per species and to the inconsistent rates of growth and survival, comparisons provided partial answers that only helped to confuse the results. Therefore, no statistical tests or results are presented here. All descriptions utilized dissecting microscopes (10-60X). An Olympus image analysis system (CUE II) was employed for the morphometric measurements. illustrations conform to the guidelines established by Sumida et al. (I 984) and descriptive terminology and methods follow Moser et al. (I 977), Moser and Ahlstrom (I 978), Richardson and Laroche (1979), and Kendall and Lenarz (1987). 1 0

RESULTS

Descriptions of Larvae

All of the species described had several common characters at birth which are shared with all Sebastes species described to date (Morris, 1956; Moser, 1967, 1972; Waldron, 1968; Westrheim, 1975; Moser et al., 1977; Moser and Ahlstrom, 1978; Richardson and Laroche, 1979; Laroche and Richardson, 1980,1981; Moser and Butler, 1981,1987; Moser, et al. 1985; Stahl-Johnson, 1985; Wold, 1990): (1) pigmented eyes and functional mouths; (2) an undifferentiated finfold surrounding the trunk and tail; (3) pigmentation on the dorsolateral surface of the gut (locus 11) and posterior surface of the hindgut (locus 17); and ( 4) pigmentation on the ventral surface of the trunk (Kendall and Lenarz, 1987). The extent of postanal midline pigmentation is a character which helps to separate groups of species (DeLacy et. al., 1964; Moser, 1967; Westrheim, 1975; Moser et. al., 1977). In this study, the larvae of two species (S. mystinus and S.melanops) had short ventral series of melanophores (loci 22 and 23), which are different from the longer, denser series (loci 21-23) characteristic of tl1e other three species (S. carnatus, S. atrovirens, and S. rastrelliger). In addition, these two species lacked, or had very little, pigmentation on the dorsal portion of the trunk (locus 19). This contrasted with the other three species which had complete dorsal series (loci 18 and 19). All broods, unless otherwise indicated, were born during night and twilight hours. 1 1

Sebastes mystinus

Larvae were obtained from three females during January and February 1989. Two broods were born having both fully absorbed and large (0.6 mrn mean diameter) yolk sacs. The latter had higher mortalities and, due to the buoyancy of the yolk sacs and oil globules, were often seen at the surface swimming on their sides and occasionally upside down. Larvae from these two broods lived 10 and 11 days, respectively. A third female gave birth to full term larvae, some of which survived 31 days. Some larvae from this brood showed signs of starvation such as dorsoventrally compressed heads, "duck billed"jaws, and reduced body depth (Figure 5). At 28 days the marine copepod Tigriopus was introduced into the diet of this brood to supplement the rotifer-brine shrimp nauplii-wild plankton diet. This copepod proved to be an unsuitable food item due to its large size and to its aggregation on the sides of the tanks where the majority of the larvae did not congregate. The larvae ranged in size from 3.2 mrn at birth to 5.5 mrn at day 31. The mean SLat birth was 3.8 mrn (SD = 0.2), with a mean of 4.7 mm (SD = 0.4) at age 31 (Figure 6). The later stages were characterized by pigmented pectoral fins that extended to the back of the anus by the time a 5.0 mm mean length (day 21) was reached (Figure 7). This species had the largest pectoral fms of any of the species described here. These larvae (Table 1; Figure 7) were born with pigmentation patterns similar to those of S. melanops (Figure 13). At birth all larvae had pigmentation on the dorsal surface of the gut (locus 11) and anus (loci 16 and l7); the short ventral midline series (loci 22 and 23) was composed of relatively few melanophores (mean number= 15.4 ). By a mean length of 5 mm (day 21), the majority of larvae had developed pigmentation on the lower 1 2 jaw (locus 1), forebrain (locus 4), nape (locus 6; Figure 8), otic capsule (locus 31), and cleithral symphysis (locus 28). Melanophores were also beginning to appear posteriorly on the dorsal surface of the trunk (locus 19; Figure 9) and on the margin and blade of the pectoral fins (loci 8 and 9). Head spines were first observed on larvae with a 4.7 mrn mean length at 31 days: the first and second posterior preoperculars, and pterotic spines were present (Figure 7c).

Sebastes carnatus

Three field-caught pregnant S. carnatus gave birth in the rearing tanks during March, April, and May 1989. One of these released larvae with large yolk sacs (0.6 mrn mean diameter) and large quantities of oil. First feeding of these larvae was observed after five days (4.0 mrn mean SL), by which time the yolk sacs were depleted. Some of the larvae lived for 28 days. The other two females gave birth to full term larvae that lived five and eight days. Leeches (Hirudinoidea: probably Piscicola, Malmiana, or Ostreobdella spp.) were found on three larvae from the longest surviving brood at 23 days. These leeches were attached (one per larva) on the left side of the head and detached themselves when the larvae were disturbed. The occurrence of this parasite probably resulted from the use of an infected female as brood stock. The range in length of these larvae was 3.1 mrn at birth to 6.2 mm at 28 days. The mean SL of yolksac larvae was 3.4 mm (SD = 0.1) at birth and 5.8 mm (SD = 0.2) at 28 days (Figure 6). The full term larvae were 4.4 mrn (SD "'0.3) mean SLat birth. The pectoral fin reached to just in front of the anus by the time a mean length of 4.3 mrn (day 13) was attained (Figure 10). 1 3

The yolksac larvae showed little pigmentation (Table 2; Figure lOa). However, complete ventral (loci 21-23) and dorsal midline (loci 18 and 19) pigment series were present (Figure 9) along with a few melanophores scattered laterally on the yolksac and a prominent anterio-lateral trunk melanophore (locus 27). Melanophores on the dorsal surface of the gut (locus 11), anus (loci 16 and 17), and the anterio- and posterio-dorsal aspect of the trunk (loci 18 and 19) were also quite frequent. All individuals in the 5.0-5.4 mm size class (day 13) had pigments on the jaw (locus 1), forebrain (locus 4), nape (locus 6; Figure 8), anus (locus 16), and ventrally on the gut (locus 10). The 6.0-6.4 mm size class (day 25) showed pigment on the maxilla (locus 2), base and border of the pectoral fins (loci 7 and 9), cleithrum (locus 28), and otic capsule (locus 31; Figure lOc). Spination was first observed on larvae of 6.1 mm mean length (25 days) with the appearance of the first and second posterior preoperculars, pterotic, and parietal spines (Figure lOc).

Sebastes atrovirens

Larvae of S. atrovirens were obtained from two females removed from the MBA Kelp Tank in May 1988 and June 1989. One female spawned naturally in the rearing tank, the other gave birth on capture to larvae and unfertilized eggs (0.8 mm mean diameter). The newborn larvae from both broods were of comparable sizes and developmental stages. Some larvae from the full term brood lived eight days; some from the mixed brood lived for 30 days.

The range of larval length was between 4.0 mm at birth and 6.5 mm at day 30. The mean length at birth was 4.3 mm (SD = 0.2) with a mean of 5.3 j 4 mm (SD = 0.6) at 30 days (Figure 6). The pectoral fms extended to just in front of the anus by the time a mean length of 4.5 mm (day 10) was reached (Figure 11). At birth, S. atrovirens larvae (Table 3; Figure 11) were similar to S. carnatus with pigmentation on the ventral and dorsal aspects of the gut (loci 10 and 11 ), the anus (locus 16), posterior aspect of the hindgut (locus 17), posterior dorsal midline of the trunk (locus 19), and a complete ventral midline series (loci 21-23). With development, increasing proportions of larvae showed pigmentation on the lower jaw (locus 1), forebrain (locus 4), nape (locus 6; Figure 8), anterior dorsal midline (locus 18; Figure 9), and otic capsule (locus 31). With continued growth, new pigmentation appeared on the posterior and posterio-ventral surfaces of the gut (loci 12 and 13), cleithrum (locus 28), and lower jaw (locus 29). The pigmentation patterns of S. atrovirens and S. carnatus remained extremely similar throughout all stages (seeS. carnatus description for explanation of differences). No head spine development was apparent during the 28 days (Figure 11).

Sebastes rastrelliger

One femaleS. rastrelliger removed from the "MBA Kelp Tank in mid­ February 1989 gave birth on capture, but the larvae appeared to be full term since no yolk sacs or oil globules were present. Some of these larvae lived up to 56 days. Larvae ranged from 4.3 mm at birth to 7.7 mm at day 52. The mean length at birth was 4.6 mm (SD = 0.1) and at day 56 was 6.6 mm (SD = 0.3; Figure 6). The pectoral fins in this species were smaller than the other species, reaching to -75% of the gut length (Figure 12). 1 5

This was the most highly pigmented of the five species described (Table 4; Figure 12). The newborn larvae had completely pigmented dorsal and ventral midline series and pigmentation on the forebrain and nape (loci 4 and 6). Melanophore numbers at these loci increased from a mean of 4.6 and 3.0 at birth to 20.0 and 18.7 at 7.5-7.9 mm (Figure 8). Melanophores on the jaw, maxilla, and posterior half of the jaw (loci 1-3), were sometimes present at birth, and were present on all larvae by the time a 6.0-6.4 mm length was reached (Table 4). Melanophore density in the anterior dorsal midline series (locus 18) increased from 3.6 to 15.7. The posterior dorsal midline series (locus 19) also increased, going from a mean of 44 to> 44 at 7.5-7.9 mm (Figure 9). Due to the dense arrangement of melanophores, it was not possible to quantify their exact number at this locus. Distinctive posterio­ lateral trunk pigment (loci 24 and 33) was present on most 5.0-5.4 mm larvae and present in all6.5-6.9 mm larvae. This pigmentation was also seen on similar-sized S. carnatus larvae. Spination was present on larvae of 6.3 mm mean SL at 29 days: the parietal, pterotic, first and second anterior preoperculars, second, third, and fourth posterior preoperculars, upper opercular, and postocular spines were present (Figure 12c). By 7.4 mm mean SLat 34 days the parietal, pterotic, first, second, and third anterior preoperculars, second, third, and fourth posterior preoperculars, upper opercular, and postocular spines were observed (Figure l2d). Hypural elements were developed on the 7.4 mm (34 day old) larvae. 1 6

Sebastes melanops

One field-captured female gave birth in February 1989 to larvae which were premature, presumably due to capture-related stress. The larvae were kept alive at the Telonicher Marine Laboratory for 22 days. Although they were offered food, no growth was observed once the yolk was consumed and they demonstrated signs of starvation similar to some S. mystinus larvae (Figure 5). As a consequence only 1-3 day old preflexion stage larvae were described here. The larval sizes ranged from 3.4 mm at birth to 4.1 mm at day 3. The mean larval length at birth was 3.6 mm (SD = 0.1), with a mean of 4.0 mm (SD = 0.1) at day 3 (Figure 6). This species had relatively small and unpigmented pectoral fins, characters that helped to differentiate it from S. mystinus. Of all the species described in this study S. melanops was the least pigmented for the size range described (Table 5; Figure 13). Along with S. mystinus these were the only early larvae lacking dorsal midline pigment (loci 18 and 19; Figure 9), and also having short ventral midline series (loci 22 and 23) composed of relatively few melanophores for the 3.5-4.4 mm size range. All S. melanops newborn larvae had pigmentation on the ventral, and dorsal surfaces of the gut (loci 10 and 11 ), the posterior surface of the hindgut (locus 17), the finfold between the gut and anus (locus 14), and the posterio-ventral surface of the notochord (locus 25). Only a small proportion of these larvae had pigment on the anus (locus 16), and the lower lobe of the caudal fm (locus 26). By day 3 pigmentation at the same loci persisted, and the frequency of occurrence of melanophores on the anus (locus 16) and the anterio-lateral section of the trunk (locus 27) increased (Table 5). 1 7

Species Comparisons a. Morphometries

Sebastes mystinus was characterized by having relatively longer snouts (SNL/HL), and deeper bodies (BD/SL) than the other species at similar sizes. In addition, this species showed somewhat larger heads (HL/SL) in relation to body length (Table 6-10). Sebastes carnatus larvae appeared to have longer guts (SA/SL) than the other species at all sizes exceptS. atrovirens at the 4.5-5.4 mm size range. It also had relatively longer snouts (SNL/HL) and deeper bodies (BD/SL) than S. atrovirens and S. rastrelliger in the 4.0-6.4 and 5.0-6.4 mm ranges (Tables 7-9). Sebastes atrovirens had somewhat longer guts (SA/SL), compared to the other species exceptS. carnatus of 4.5-5.4 mm. More notably this species had fairly large eyes (ED/HL) that may help to distinguish it from the very similar S. carnatus larvae (Tables 8 and 9). Sebastes rastrelliger, along with S. mystinus, had relatively shorter guts (SA/SL), and deeper bodies (BD/SL) compared with the other species. It also had smaller eyes (ED?HL) than S. carnatus and S. mystinus (Tables 6, 7 and 9). Sebastes melanops can be differentiated quite readily by pigmentation characters from all species exceptS. mystinus, and was therefore compared morphometrically to the latter. The morphometric proportions did not appear to be very different from those of S. mystinus, a closely related species. Sebastes melanops appeared to have larger eyes (ED/HL) than all other species for the 3.5-4.4 mm size class (Table 10). It also seemed to have longer guts (SA/SL) than S. mystinus (Table 6). 1 8 b. Pigmentation

The pigment patterns which help to differentiateS. mystinus from the other species described are: (1) a short row of ventral midline melanophore series (loci 22 and 23); (2) absence of dorsal midline pigment on the early larvae, and presence of relatively sparse posterio-dorsal midline pigment (locus 19; Figure 9) in larger specimens; and (3) presence of pigment on the pectoral fins (loci 8 and 9) with melanophores scattered over the fin rays and concentrated on the margin (Figure 7). Sebastes carnatus (Figure 10) and S. atrovirens (Figure 11) can be differentiated from S. mystinus (Figure 7) and S. melanops (Figure 13) by: (1) presence of a long ventral midline melanophore series (loci 21-23); and (2) greater number of melanophores on the dorsal midline (loci 18 and 19; Figure 9). They can be differentiated from S. rastrelliger (Figure 12) by the overall higher pigmentation on the latter. Sebastes carnatus can be differentiated from similarly pigmented S. atrovirens by the following characters: (1) frequent presence (up to 0.9) of anterio-lateral trunk pigment (locus 27; Table 3); (2) presence of melanophores on the base and anterior margin of the pectoral fins (loci 7 and 9); and (3) presence of melanophores on the midupper and midlower jaw (loci 2 and 3; Table 3; Figure 10). Sebastes rastrelliger can be differentiated by the following patterns: (1) heavily pigmented head (loci 1-4 and 6; Figures 8 and 12) with presence of melanophores on the nares (locus 32); (2) complete, heavily pigmented dorsal midline series (loci 18 and 19; Figure 9) from birth; and (3) external and internal pigmentation on the posterio-lateral portion of the trunk (loci 24 and 33) in larvae >5.0 mm (Figure 12; Table 4). TI1e extent and I 9 concentration of ventral midline pigment (loci 21-23) is similar to that of S. carnatus and S. atrovirens. Sebastes melanops had low melanophore densities with the following patterns differentiating it from the other species: (1) a short, thin row of ventral midline melanophores , similar to that of S. mystinus but narrower; (2) lack of pigmentation on the forehead (locus 4 ), nape (locus 6; Figure 7), and dorsal midline (loci 18 and 19; Figure 9); and (3) presence of melanophores on the caudal fin (loci 25 and occasionally 26; Table 5; Figure 13). 20

DISCUSSION

The traditional method of working backwards from field caught identifiable juveniles to newborn larvae may not be practical for all species of Sebastes since many field caught Sebastes larvae smaller than 8-10 mm cannot be recognized with confidence (Kendall and Lenarz, 1987). The very early stages of S. carnatus and S. atrovirens (this study), S. chrysomelas (Wold, 1990), S. auriculatus and S. caurinus (Stahl-Johnson, 1985) show great similarities and are difficult to differentiate from each other. Thus, the correct description of these species from the series approach may have not been possible. Of the 51 species of eastern Pacific Sebastes larvae previously described, only nine have been made from reared specimens (Moser and Butler, 1981, 1987; Stahl-Johnson, 1985; Wold, 1990). This is mainly due to the difficulty of rearing the larvae past the yolk sac and to the juvenile stage. Such difficulties arise from the relatively small size, early developmental level at birth, feeding problems and very importantly, slow larval growth rates shown for the eastern Pacific species (Moser and Butler, 1987). One major concern in rearing Sebastes is the need for healthy full-term larvae (Westrheim, 1975). Most information on the early larval stages of rockfishes has come from descriptions of preextrusion larvae. This presents problems because pigment patterns are not well established in yolksac Sebastes larvae, making these descriptions unreliable for the identification of more advanced stages (Westrheim, 1975). This problem is alleviated for nearshore Sebastes species with the use of the SCUBA-aided capture techniques employed in this study. It may also be possible to capture healthy females of deeper dwelling species for which information on early stages is 21 lacking (e. g. [S.fiavidus] and chilipepper [S. goodei]), by bringing hook and line caught fish to 20-30 m depths (where the largest decompression change occurs) and allowing SCUBA divers to deflate the gas bladders, and if needed, to bag the fish for further acclimatization at depth. The use of divers allows for close monitoring of the fish's condition, and faster and less stressful capture reducing the chance of abortion or premature spawning. In the case of shallow living species, this technique allows for species and condition-selective fishing to occur. Blaxter (1970), Johannes (1978), and Hunter (1981) suggested that parental behavior affects the vulnerability of pelagic eggs and larvae to predation. Time, location, and density of egg and larval release may reduce the effects of predation on these stages. Species in Sebastes seem to follow this pattern with natural spawnings occurring at twilight and night, reducing the vulnerability of larvae to diurnal predators. In addition, they spawn in batches, presumably reducing the chance of predation by lowering the densities of larvae released at a time. Also, since larval food has a patchy distribution and larvae are usually able to stay with these patches once found (Hunter and Thomas, 1974; Lasker, 1975; Methot and Kramer, 1979), batch spawning may increase the chances of larvae to encounter and remain with the food patch until more favorable conditions are found. Time of the year also seems to play a role in the time of parturition, since most Sebastes species spawn during late winter-spring season, associated with upwelling along the west coast of North America. The months of parturition observed for three of the five species (S. mystinus, S.carnatus, and S. melanops) were similar to those reported by Wyllie­ Echeverria (1985). There is no published information on parturition periods of S. atrovirens or S. rastrelliger. Sebastes species seem to have two major 22 seasons (winter and spring-summer) of larval extrusion (Phillips, 1964). Sebastes melanops, S. mystinus, and S. rastrelliger gave birth in the winter season (November-March), and S. atrovirens in the spring-summer season (April-July). Sebastes camatus overlapped both seasons with March-May parturitions. On four occasions, females released either larvae of different developmental stages (S. mystinus), larvae and oil (S. carnatus), or larvae and unfertilized eggs (S. atrovirens). There may be two explanations for these notable births. (Siokawa and Tsukahara, 1961) in the western Pacific, and S. paucispinis (Moser, 1967), S. ovalis and S. constellatus (MacGregor, 1970) in the eastern Pacific, are capable of multiple births within a reproductive season. This may also apply to S. mystinus and possibly the other species. A second theory is that the eggs or oil may be used for nutrition by the embryos (Boehlert and Y oklavich, 1984; Boehlert et al., 1987) or, as observed in anchovies (Hunter and Kimbrell, 1980), by the free swimming larvae. Over the two rearing seasons, a number of changes were made in the rearing techniques to increase survival,. The most important of these appeared to be the increase in number and variety of food items and the reduction of larval densities in rearing tanks. High densities of food organisms have been shown to be critical for successful larval Sebastes rearing (Stahl-Johnson, 1985). The quality of the food is also of concern, since larvae raised on mono diets are susceptible to high mortalities and deformed or incomplete development. This was ameliorated in the present study by inclusion of newly hatched, supplemented brine shrimp (Artemia) nauplii, enriched rotifers, and daily caught wild plankton. Low larval densities in the rearing tanks proved to be essential to the survival and growth 23 of the species treated in the study. Although Stahl-Johnson (1984) recommended a density ::::; 2larvae/L for rearing Sebastes larvae, the densities used in this study were higher (between 5-10 larvae/L). It is thought that rockfish larvae are visual feeders, and therefore do not feed at night. The introduction of permanent lighting may have improved survival and growth as observed by Boehlert (1981) while rearing S. diploproa. Temperature has been been shown to greatly affect growth of larval and juvenile rockfish (Howell, 1980; Boehlert, 1981; Boehlert and Yoklavich, 1983). Cold temperatures (9-12 °C) in Monterey Bay during the study period may have had an impact on the already slow growth rates of the larvae. For future studies it may be beneficial to increase the water temperature of the tanks to 15 °C as done for juvenileS. diploproa (Boehlert, 1981), or higher as done for S. melanops (Boehlert and Yoklavich, 1983) to allow faster growth and therefore more complete developmental series. Only the larvae of S. rastrelliger were maintained past the preflexion stage. These larvae lived for up to 56 days and went through flexion but did not reach postflexion. Although only one of the species survived through flexion, the data presented here are still very useful since small, preflexion forms (e. g.,::::; 7 mm) generally dominate (> 90% ) the Sebastes larvae caught off California by net sampling operations (Moser and Butler, 1987). To describe the Sebastes larvae it was necessary to choose a character, either age or size, to allow an evaluation of variability, and a determination of useful specific identifying characters. As in most studies of Sebastes larvae, size was used instead of age. This was done because (a) grouping by age is a slow and expensive process in field studies and (b) size rather than age appears to be a better indicator of development, although there is variability in both. Policansky (1982) found that size (length), and not age, was the single 24

most important determinant in the timing of metamorphosis of the starry flounder, Platichthys stellatus. He further stated (1983) that size at first metamorphosis was a characteristic commonly expressed throughout the kingdom. Morphometric measurements did not yield any differences useful to positively identify a species. Sebastes mystinus had relatively larger pectoral fins than the other species, but due to the difficulty in measuring the unpigmented pectoral fms of the other species it was not possible to statistically compare them. Sebastes melanops had larger eyes, and S. mystinus and S. rastrelliger had shorter guts, relative to the other species. Sebastes mystinus generally had larger heads and longer snouts than the other species at all sizes, while S. atrovirens in the 4.5-6.4 mm size range had larger eyes. With the exception of S. camatus and S. atrovirens, most species can be told apart by the difference in pigmentation. Small differences in morphometries may be used to distinguish these two species; S. carnatus had deeper bodies and smaller eyes than S. atrovirens. Two distinct pigmentation patterns can be discerned among the larvae described here: (1) a short row of ventral melanophores and no, or very little, posterio-dorsal pigmentation (S. mystinus and S. melanops); and (2) complete ventral pigmentation (e. g., all three loci pigmented) and anterior and posterio-dorsal melanophores (S. carnatus, S. atrovirens, and S. rastrelliger). Sebastes mystinus, although similar to S. melanops, had pigmentation on the head and on the margin and blade of the pectoral fins. Sebastes carnatus and S. atrovirens had similar melanophore patterns in the very early stages, with S. camatus differing in the later stages by addition of melanophores to the base and margin of the pectoral fin. Sebastes rastrelliger · was the most pigmented species with a greater number of pigmented loci and

''····'"" . 25 more melanophores per locus than any of the other species. This species was unique here by having melanophores on the nares. The internal posterio­ lateral pigmentation on the trunk of S. rastrelliger and S. carnatus is also present inS. caurinus and S. auriculatus (Stahl-Johnson, 1985) and may commonly occur in older larvae; its absence on the other species described here could be due to their shorter rearing times and growth. Sebastes melanops had the least amount of pigmentation, with a characteristic short ventral series and no head or dorsal melanophore series. Sebastes melanops larvae from Oregon described by Kendall (1989) showed head pigmentation not present on the larvae in this study. This may be due to clinal variations from Oregon to California, or to intraspecific differences that were not observed in either study due to the small sample sizes. It is relatively easy to identify three (S. mystinus, S. rastrelliger, and S. melanops) of the five species described here. However, S. carnatus and S. atrovirens are very similar and pigmentation alone will not be enough to identify all the stages of either species. Here it is necessary to utilize morphometric measurements which, although of limited use, may help verify an identification. Parturition times and range information are also very important to discern which species may be in the plankton at a certain time and place. Other approaches to the identification of Sebastes larvae include capture and rearing of planktonic larvae (Kendall, in press) and electrophoretic techniques (Seeb and Kendall, 1990). However, these methods, have limitations. Capture of viable Sebastes larvae is difficult since they are extremely delicate and are not very accessible (e. g., most individuals are collected at ca. 40 m depths; Ahlstrom, 1959). In addition, a typical plankton tow takes about 20 minutes (Smith and Richardson, 1977) while autolytic 26 tissue decomposition occurs within two to three minutes after sampling (O'Connell, 1980; Setzler-Hamilton et al., 1987) possibly affecting electrophoretic results. Some investigators (Morris, 1956; DeLacy et al., 1964; Moser, 1967; Efremenko and Lisovenko, 1970; Westrheim, 1975) have serious doubts about the ability to identify field caught Sebastes larvae due to the often seen small interspecific differences and large intraspecific variabilities. However, this is not the case for some already described species, and may not be the case for most species. There is a need for researchers to describe in more detail the degree of variability of characters commonly used for species identification (e. g., pigmentation and spination). Due to the great similarities between some species and to possible sampling related morphological changes (Theilacker, 1980) it may also be necessary to modify sampling techniques if species identification is desired for larvae in this genus. Some sampling modifications may include shorter towing times, smaller nets, and faster processing both before and after fixation. In addition, it is important to utilize all information concerning ranges and reproductive periods for all species to help make, in the worst of cases, an educated guess. 27 LITERATURE CITED

Ahlstrom, E. H. 1959. Vertical distribution of pelagic fish eggs and laiVae off California, and Baja California. U.S. Fish. Wildl. SeiV., Fish. Bull.161: 107-146.

Bainbridge, V., and B.J. McKay. 1968. The feeding of cod and redfish. Spec. Pub!. of Int. Comm. N.W. Atlantic Fish. No.7: 187-217.

Barsukov, V. V. 1981. A brief review of the subfamily Sebastinae. J. Ichthyol. 21: 1-26.

Blaxter, J. H. S. 1970. Light. In: Marine Ecology, a comprehensive, integrated treatise on life in the oceans and coastal waters, Vol. 1. Environmental Factors, (ed. 0. Kinne), pp. 213-320, Wiley­ Interscience, New York.

Boehlert, G. W. 1981. The effect of photoperiod and temperature on laboratory growth of juvenile , and a comparison with growth in the field. Fish. BulL 79(4): 789-794.

Boehlert, G. W., and M. M. Yoklavich. 1983. Effects of temperature, ration, and fish size on growth of juvenile black rockfish, Sebastes melanops. Environ. Biol. Fish. 8: 17-28.

Boehlert, G. W., and M. M. Yoklavich. 1984. Reproduction, embryonic energetics, and the maternal-fetal relationship in the viviparous genus Sebastes (Pisces: Scorpaenidae). Bioi. Bull. 167: 354-370.

Boehlert, G. W., M. Kusakari, and J. Yameda. 1987. Estimating the cost of viviparity in the genus Sebastes, p. 99-128. In: Proc. Int. Rockfish Symp., University of Alaska, Alaska Sea Grant Report No. 87-2, Anchorage, Alaska.

DeLacy, A. C., C. R. Hitz, and R. L. Dryfoos. 1964. Maturation, gestation, and birth of rockfish (Sebastodes) from Washington and adjacent waters. Wash. Dep. Fish., Fish. Res. Pap. 2(3): 51-67.

Efremenko, V. M., and L. A. Lisovenko. 1970. Morphological features of intraovarian and pelagic laiVae of some Sebastodes species inhabiting the gulf of Alaska. In: Moiseev, P. A. (ed.), Soviet fisheries investigations in the northeastern Pacific, Part. V, p. 267- 28 286. Izv. Tikhookean. Nauchno-Issled. Inst. Morsk. Rybn. Khoz. Okeanogr. (VNIRO) 70. [Trans!. from Russ.: avail. U.S. De. Commer., N atl. Tech. Inf. Serv. Springfield, VA 22161, as TTII- 50127.

Eschmeyer, W. N., E. S. Herald, and H. Hammann. 1983. A Field Guide to Pacific Coast Fishes of North America. Houghton Mifflin Co., Boston, 336 p.

Fujita, S. 1957. On the larval stages of a scorpaenid fish, nigricans (Schmidt). Jap. J. Ichthyology. 6: 91-93. (In Japanese with English summary).

Fujita, S. 1958. On the egg development and larval stages of a viviparous scorpaenid fish, Sebastes oblongus Gunther. Bull. Jap. Soc. Sci. Fish. 24: 475-479. (In Japanese with English summary).

Guillard, R. R. L. 1975. Culture of phytoplankton for feeding marine invertebrates., p. 29-60. In: Culture of Marine Invertebrate . Smith, W. L. and M. H. Chanley (eds), Plenum Press, New York and London.

Gunderson, D. R. 1977. Population biology of Pacific ocean perch, Sebastes alutus, in the Washington-Queen Charlotte Sound, Oregon, and their response to fishing. Fish. BulL 75(2): 369-403.

Gunderson, D. R., and W. H. Lenarz. 1980. Cooperative survey of rockfish and whiting resources off California, Washington, and Oregon, 1977. Marine Fish. Review 42: 2.

Haldorson, L. 1987. Introduction, p.1-4. In: Proc. Int. Rockfish Symp., University of Alaska, Alaska Sea Grant Report No. 87-2, Anchorage, Alaska.

Hempel, G. 1973. (ed.) Fish eggs and larval studies. (Contributions to a manual). FAO Fish. Tech. Pap., (122): 82 pp.

Hoshiai, G. 1977. Larvae and juveniles of the scorpaenid fish, Sebastes schlegeli. Jap. J. Ichthyology 24: 35-42. (In Japanese with English summary). 29 Howell, W. H. 1980. Temperature effects on growth and yolk utilization in yellowtail flounder, Limanda ferruginea, yolk sac larvae. Fish. Bull. 78(3): 731-739.

Hughes, J. T., R. A. Shleser, and G. Tchobanoglous. 1974. A rearing tank for lobster larvae and other aquatic species. The Progressive Fish­ Culturist 36(3): 129-132.

Hunter, J. R. 1981. The feeding ecology of marine fish larvae. In: Fish behavior and its use in the capture and culture offishes (eds. J. E. Bardach, J. J. Magnuson, R. C. May, and J. M. Reinhart), ICLARM Conf. Proc. 5, Intemtl. Center for Living Aquatic Resources Mgmt., Manila. p. 287-330.

Hunter, J. R., and C. A. Kimbrell. 1980. Egg cannibalism in the northern anchovy, Engraulis mordax. Fish. Bull. 78(3): 811-816.

Hunter, J. R. and G. L. Thomas. 1974. Effect of prey distribution and density on the searching and feeding behavior of larval anchovy Engraulis mordax Girard, pp. 559~574. In: The Early Life History of Fish, (ed. J. H. S. Blaxter), Springer-Verlag, Berlin.

Johannes, R. E. 1978. Reproductive strategies of coastal marine fishes in the tropics. Env. Biol. Fish. 3:65-84.

Kendall, A. W., Jr. in press. Systematics and identification of larvae and juveniles of the genus Sebastes. Environ. Bioi. Fish.

Kendall, A. W. Jr. 1989. Additions to knowledge of Sebastes larvae through recent rearings. Northwest and Alaska Fisheries Center Processed Report 89-XX; XXpp.

Kendall, A. W., Jr., and W. H. Lenarz. 1987. Status of early life history studies of northeast Pacific rockfishes, p. 99-128. In: Proc. Int. Rockfish Symp., University of Alaska, Alaska Sea Grant Report No. 87-2, Anchorage, Alaska.

Kusakari, M., Y. Mori, and K. Kudo. 1977. Studies on the breeding habit of a rockfish, Sebastes schlegeli (Hilgendorf) 2. On the breeding behavior of pregnant fish and just spawned larvae. Hokkaido Fisheries Experimental Station Monthly Report. Vol. 34(10) Supplement. 11 p. 30 Laroche, W. A., and S. L. Richardson. 1980. Development and occurrence of larvae and juveniles of the rockfishes Sebastes jlavidus and Sebastes melanops (Scorpaenidae) off Oregon. Fish. Bull., U. S. 79: 901-924.

Laroche, W. A., and S. L. Richardson. 1981. Development of larvae and juveniles of the rockfishes Sebastes entomelas and S. zacentrus (Family Scorpaenidae) and occurrence off Oregon, with notes on head spines of S. mystinus, S. jlavidus, and S. melanops. Fish. Bull. 79(2): 231-257.

Lasker, R. 1975. Field criteria for survival of anchovy larvae: lhe relation between inshore chlorophyll maximum layers and successful first feeding. U. S. Fish. Bull. 73: 453462.

Lenarz, W. H. 1987. A history of California rockfish fisheries, p.35-41. In: Proc. Int. Rockfish Symp., University of Alaska, Alaska Sea Grant Report No. 87-2, Anchorage, Alaska.

Love, M.S., and W. V. Westphal. 1981. Growth, reproduction, and food habits of olive rockfish, , off central California. Fish. Bull. 75(2): 412-416.

MacGregor, J. S. 1970. Fecundity, multiple spawning, and description of gonads in Sebastodes. U.S. Fish Wildl. Serv ., Spec. Sci. Rep. Fish. 596, 12 pp.

Melhot, R. D., and D. Kramer. 1979. Growth of northern anchovy, Engraulis mordax, larvae in lhe sea. U.S. Fish. Bull. 77: 413-423.

Morris, R. W. 1956. Early larvae offour species of rockfish, Sebastodes. Calif. Fish and Game 42: 149-153 ..

Moser, H.G. 1967. Reproduction and development of Sebastodes paucispinis and comparison wilh olher rockfishes off southern California. Copeia: 773-797.

Moser, H. G. 1972. Development and geographic distribution of the rockfish, Sebastes macdonaldi (Eigenmann and Beeson, 1893), family Scorpaenidae, off southern California and Baja California. Fish. Bull. 70(3) 941-958. 31 Moser, H. G., and E. H. Ahlstrom. 1978. Larvae and pelagic juveniles of blackgill rockfish, Sebastes melanostomus, taken in mid water trawls off southern California and Baja California. J. Fish. Res. Bd. Can. 35: 981-996.

Moser, H. G., E. H. Ahlstrom, and E. M. Sandknop. 1977. Guide to the identification of scorpionfish larvae (family Scorpaenidae) in the eastern Pacific with comparative notes on species of Sebastes and from other oceans. NOAA Tech.Rep. NMFS Circ. 402. 71 p.

Moser, H. G., and J. L. Butler 1981. Description of reared larvae and early juveniles of the calico rockfish, . Calif. Coop. Ocean. Fish. Invest. Rep. 22: 88-95.

Moser, H. G., and J. L. Butler. 1987. Descriptions of reared larvae of six species of Sebastes, p. 19-29 In: , Proceedings of a workshop. Lenarz W. H. and D. R. Gunderson (eds), NOAA Technical Report NMFS 48: 57 pp.

Moser, H. G., E. M. Sandknop, and D. A. Ambrose. 1985. Larvae and juveniles of Aurora rockfish, , from off California and Baja California, p. 65-76. ln: Descriptions of early life history stages of selected fishes. Kendall, A. W., Jr. and J. B. Marliave (eds), Can. Tech. Rep. Fish. Aquat. Sci. 1359.

O'Connell, C. P. 1980. Percentage of starving northern anchovy, Engraulis mordax, larvae in the sea as estimated by histological methods. Fish. Bull. 78: 475-489.

Phillips, J. B. 1964. Life history studies of ten species of rockfish (genus Sebastodes). Calif. Dep. Fish Game, Fish. Bull. 126, 70p.

Policansky, D. 1982. Influence of age, size, and temperature on metamorphosis in the starry flounder, Platichthys stellatus. Can. J. Fish. Aquat. Sci. 39: 514-517.

Policansky, D. 1983. Size, age and demography of metamorphosis and sexual maturation in fishes. Am. Zool. 23: 57-63.

Richardson, S. L., and W. A. Laroche. 1979. Development and occurrence of larvae and juveniles of the rockfishes Sebastes 32 crameri, Sebastes pinniger, and (Family Scorpaenidae) off Oregon. Fish. Bull. 77(1): 1-41.

Seeb, L. W., and A. W. Kendall, Jr. 1990. Allozyme and DNA polymorphisms: New approaches to the identification of larvae and of genus Sebastes. In Press.

Setzler-Hamilton, D.A. Wright, F.D. Martin, C.V. Millsaps, and S.I. Whitlow. 1987. Analysis of nutritional condition and its use in predicting striped bass recruitment: field studies. Amer. Fish. Soc. Syrup. 2: 115-128.

Siokawa, T., and H. Tsukahara. 1961. Studies on habits of coastal fishes in the Amakusa Islands. Part 1. Early life history of the purple rockfish, Sebastes pachycephalus pachycephalus Temmick et Schleger. Rec. Oceanogr. Works Jap., Spec. No.5: 123-127

Smith, P. E., and S. L. Richardson. 1977. Standard techniques for pelagic fish egg and larva surveys. FAO Fish. Tech. Paper No. 175, Rome.

Stahl-Johnson, K. L. 1984. Rearing and development of larval Sebastes caurinus () and S. auriculatus () from the northeastern Pacific. M. Sc. Thesis, Univ. Washington, Seattle. 217 pp.

Stahl-Johnson, K. L. 1985. Descriptive characteristics ofreared Sebastes caurinus and S. auriculatus larvae, p. 65-76 In: Descriptions of early life history stages of selected fishes. Kendall, A. W. Jr., and J. B. Marliave (eds). Can. Tech. Rep. Fish. Aquat. Sci. 1359.

Sumida, B. Y., B. B. Washington, and W. A. Laroche. 1984. Illustrating fish eggs and larvae, p. 33-35. In: Ontogeny and Systematics of Fishes. Moser, H. G., W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall Jr., and S. L. Richardson (eds). Spec. Publ. No. 1, Amer. Soc. Ichthyol. Herpetol.

Takai, T., and T. Fukunaga. 1971. The life history of an ovoviviparous scorpaenoid fish, Sebastes longispinis (Matsubara). 1. Egg and larval stages. J. Shimonoseki Univ. Fish. 20: 91-95. (In Japanese with English summary). 33 Theilacker, G. H. 1980. Changes in body measurements of larval northern anchovy, Engraulis mordax, and other fishes due to handling and preservation. Fish. Bull. 78(3): 685-692.

Waldron, K. D. 1968. Early larvae of the , Sebastodes pinniger. J. Fish. Res. Board Can. 25: 801-803.

Westrheim, S. J. 1975. Reproduction, maturation, and identification of larvae of some Sebastes (Scorpaenidae) species in the northeast Pacific Ocean. J. Fish. Res. Bd. Canada. 32(12): 2399-241 L

Wold, L. 1990. A practical approach to the description and identification of Sebastes larvae. M. Sc. Thesis, Moss Landing Marine Laboratories, Moss Landing, CA. 69 pp.

Wyllie-Echeverria, T. 1985. Thirty-four species of California rockfishes: maturity and seasonality of reproduction. Fish. Bull., U. S. 85: 229- 250. SL 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 (mm) 3.0-3.4 1.0 0.0 0.0 0.0 0.0 0.8 0.0 0.0 1.0 0.3 1.0 0.3 0.0 0.8 0.3 1.0 1.0 3.5·3.9 0.2 0.0 0.0 0.1 0.0 0.1 0.0 0.0 0.1 0.4 1.0 0.0 0.0 0.2 0.1 1.0 1.0 4.0-4.4 0.6 0.0 0.0 0.4 0.0 0.4 0.0 0.4 0.6 0.6 1.0 0.1 0.0 0.3 0.0 0.9 1.0 4.5-4.9 1.0 0.0 0.0 1.0 0.2 0.8 0.0 0.7 1.0 0.8 1.0 0.3 0.2 0.5 0.3 0.8 1.0 5.0-5.4 1.0 0.0 0.0 1.0 0.2 1.0 0.0 1.0 1.0 1.0 1.0 0.2 0.2 0.0 0.2 1.0 1.0

SL 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n (mm) 3.0-3.4 0.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.0 0.3 0.3 0.0 0.5 0.0 0.0 4 3.5-3.9 0.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 38 4.0-4.4 0.1 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.0 0.6 0.0 0.0 0.2 0.0 0.0 21 4.5-4.9 0.0 . 0.3 0.2 0.0 1.0 1.0 0.0 0.5 0.5 0.0 0.8 0.0 0.3 0.3 0.0 0.0 6 5.0-5.4 0.0 0.4 0.6 0.0 1.0 1.0 0.0 0.4 0.4 0.0 1.0 0.0 0.0 0.8 0.0 0.0 5 SI. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 (mm) 3.0-3.4 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.1 1.0 0.1 0.0 0.0 0.0 1.0 1.0 3.5-3.9 0.4 0.0 0.0 0.2 0.0 0.3 0.0 0.0 0.0 0.5 1.0 0.0 0.0 0.3 0.2 0.9 1.0 4.0-4.4 0.2 0.0 0.0 0.3 0.0 0.7 0.0 0.0 0.0 0.9 1.0 0.1 0.2 0.2 0.1 1.0 1.0 4.5-4.9 0.1 0.0 0.0 0.1 0.0 0.8 0.0 0.0 0.0 0.8 1.0 0.0 0.0 0.1 0.0 1.0 1.0 5.0-5.4 1.0 0.5 0.5 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.1 0.1 0.0 1.0 1.0 1.0 5.5-5.9 1.0 0.4 0.6 1.0 0.0 1.0 0.9 0.0 0.7 1.0 1.0 0.7 0.4 0.1 0.9 1.0 1.0 6.0-6.4 1.0 0.8 0.6 1.0 0.0 1.0 1.0 0.0 1.0 1.0 1.0 1.0 1.0 0.4 1.0 1.0 1.0

SL 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n (mm) 3.0-3.4 0.8 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.9 0.0 0.0 0.0 0.0 0.0 0.0 16 3.5-3.9 0.6 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.5 0.2 0.0 0.0 0.4 0.0 0.0 13 4.0-4.4 0.9 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.2 0.1 0.0 0.0 0.3 0.0 0.0 59 4.5-4.9 0.8 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.2 0.1 0.0 0.0 0.1 0.0 0.0 24 5.0-5.4 1.0 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.0 1.0 0.4 0.0 1.0 0.0 0.0 10 5.5-5.9 1.0 1.0 0.0 1.0 1.0 1.0 0.7 0.0 0.1 0.1 1.0 0.1 0.0 1.0 0.0 0.3 7 6.0-6.4 1.0 1.0 0.0 1.0 1.0 1.0 0.8 0.0 0.0 0.6 1.0 1.0 0.0 1.0 0.0 0.6 5 1-j §...... ,(1>

"d 'i:i til ,g "'(1> 0 ::t.... ::4 P> 0~· ~..... ::l SL 1 2 J 4 5 6 7 8 9 10 11 12 13 14 15 16 17 0 "' (mm) n 0...... , 4.0-4.4 0.1 0.0 0.0 0.1 0.0 0.1 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.1 0.0 1.0 1.0 -· -.....,' ~ 4.5-4.9 0.2 0.0 0.0 0.6 0.0 0.6 0.0 0.0 0.0 1.0 1.0 0.1 0.1 0.1 0.1 1.0 1.0 g- 5.0-5.4 1.0 0.0 0.0 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.3 0.3 0.0 0.0 1.0 1.0 """(/.) ..."" (1> ~ (1> l:l ...... "rj ~ ;::;;, ~ .... -!>- "';:,: ...,0' "" ..... !>'- 0 ~ n (1> SL 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n c..~· "' (mm) (1> ...... ~ 4.0·4.4 0.3 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.0 0.0 0.0 0. 0.1 0.0 0.0 32 "'n s- :::1. "d a 4.5-4.9 0.4 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.0 0.3 0.0 0.0 0.3 0.0 0.0 19 ;:t. (1> 0 5.0-5.4 1.0 1.0 0.0 1.0 1.0 1.0 0.0 0.0 0.0 0.0 0.7 0.3 0.0 0.7 0.0 0.0 3 ::l §- ':-' ,g ::r ~ "'

w en SL 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 (mm) 4.0-4.4 0.6 0.4 0.4 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.0 OA 1.0 4.5-4.9 0.3 0.2 0.3 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.1 0.1 0.4 1.0 5.0·5.4 1.0 1.0 0.8 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.2 0.0 0.0 0.0 0.7 1.0 5.5-5.9 1.0 0.9 0.9 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.3 0.8 1.0 6.0-6.4 1.0 1.0 1.0 1.0 0.0 1.0 0.0 0.0 0.0 0.9 1.0 0.0 0.1 0.0 0.3 1.0 1.0 6.5-6.9 1.0 1.0 1.0 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.4 1.0 1.0 7.0-7.4 1.0 1.0 1.0 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.3 0.0 0.0 0.3 0.5 1.0 7.5-7.9 1.0 1.0 1.0 1.0 0.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 0.0 0.0 0.7 1.0

SL 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n (mm) 4.0-4.4 1.0 1.0 0.0 1.0 1.0 1.0 0.6 0.0 0.0 0.2 0.4 0.2 0.0 0.0 0.4 0.0 5 4.5·4.9 1.0 1.0 0.0 1.0 1.0 1.0 0.6 0.0 0.0 0.3 0.7 0.3 0.0 0.3 0.3 0.0 27 5.0-5.4 1.0 1.0 0.0 1.0 1.0 1.0 0.9 0.0 0.0 0.1 1.0 0.8 0.0 0.7 0.9 0.0 25 5.5-5.9 1.0 1.0 0.0 1.0 1.0 1.0 1.0 0.0 0.0 0.0 1.0 0.8 0.0 0.9 0.9 0.8 18 6.0-6.4 1.0 1.0 0.0 1.0 1.0 1.0 0.9 0.0 0.0 0.1 1.0 0.8 0.0 0.9 0.9 0.8 15 6.5-6.9 1.0 1.0 0.0 1.0 1.0 1.0 1.0 0.0 0.0 0.0 1.0 0.9 0.0 0.9 1.0 1.0 13 7.0-7.4 1.0 1.0 0.0 1.0 1.0 1.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 1.0 1.0 1.0 6 7.5-7.9 1.0 1.0 0.0 1.0 1.0 1.0 1.0 0.0 0.0 0.0 1.0 1.0 0.0 0.7 1.0 1.0 3 SL 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 (mm) 3.5·3.9 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 1.0 0.0 0.2 1.0 4.0•4.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1.0 1.0 0.0 0.0 1.0 0.0 1.0 1.0

SL 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n (mm) 3.5·3.9 0.0 0.0 0.0 0.0 1.0 1.0 0.0 1.0 0.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 24 4.0•4.4 0.0 0.0 0.0 0.0 1.0 1.0 0.0 0.9 0.2 0.1 0.0 0.0 0.0 0.0 0.0 0.0 15 39

Table 6: Morphometric proportions of Sebastes mystinus larvae. Mean standard deviation, with the ranges in parentheses. SA/SL= snout-anus length/standard length; HL/SL =head length/standard length; SNL/HL = snout length/head length; ED/HL = eye diameter/ head length; BD/SL =body depth/ standard length. See Figure 3.

SL SA/SL HL/SL SNLIHL ED/HL BD/SL n (mm)

3.0-3.4 39.4± 1.9 24.2 ± 1.9 37.8 ± 3.9 37.8 ± 3.9 18.9 ± 1.6 4 (38.2-40.6) (21.8-26.4) (33.3-42.8) (33.3-42.8) (17.6-21.2)

3.5-3.9 34.9± 2.2 19.4±1.6 42.1 ± 5.1 37.8 ± 6.9 16.5 ± 1.3 38 (28.2-38 .8) (15.3-23.0) (33.3-57.1) (25 .0-50.0) (12.8-19.4)

4.0-4.4 34.9 ± 1.7 20.6 ± 1.7 37.2 ± 5.1 35.8 ± 3.4 15.9 ± 2.0 21 (31.7-39.5) (18.6-25.0) (25.0-45.4) (25.0-40.0) (12.1-19.5)

4.5-4.9 37.7 ± 1.4 22.6± 2.0 46.0 ± 5.4 30.5 ± 5.5 17.2 ± 1.9 6 (35.5-39.1) (20.0-26.0) (40.0-54.5) (25.0-40.0) (15.2-19.5)

5.0-5.4 37.2 ± 1.0 22.7 ± 1.0 48.3 ± 4.9 34.5 ± 1.6 17.6±1.5 5 (36.0-38.4) (21.5-24.0) (41.6-54.5) (33.3-36.3) (16.0-20.0) 40

Table 7: Morphometric proportions of Sebastes carnatus larvae [x ± SD (range)]. See Table 6 for explanation.

SL SA/SL HL/SL SNL/HL ED/HL BD/SL n (mm)

3.0-3.4 40.5 ± 2.6 15.9 ± 2.2 24.6 ±7.9 41.2 ± 5.0 9.8 ±4.5 16 (35.2-45.1) (11.7-21.2) (16.6-40.0) (33.3-50.0) (5.8-21.2

3.5-3.9 39.7 ± 1.9 17.8±2.6 34.1 ± 7.4 37.7 ± 7.9 12.4 ± 4.5 14 (37 .1-44.4) (14.2-22.2) (20.0-42.8) (28.5-60.0) (5.5-17.9)

4.0-4.4 38.8 ±2.0 18.7 ± 1.6 34.2 ± 6.0 37.0 ± 5.8 13.3 ± 5.0 59 (34.8-42.8) (15.9-23.2) (25.0-50.0) (25.0-50.0) (2.0-19.0)

4.5-4.9 36.8± 1.4 18.5 ± 1.1 32.9 ± 5.1 35.3±7.1 14.5 ± 1.4 24 (34.7-40.0) (17.0-20.0) (22.2-37.5) (22.2-50.0) (12.7-17.7)

5.0-5.4 37.8± 1.5 20.5 ± 1.0 33.6 ±4.1 34.6 ± 4.3 17.5 ± 6.3 10 (35.8-40.7) (19.2-22.2) (27 .2-40.0) (27 .2-40.0) (14.0-35.1)

5.5-5.9 42.4± 1.8 24.2± 1.7 33.2± 3.9 33.3 ± 2.4 20.0 ± 1.4 7 (39.6-44.8) (22.0-27.1) (26.6-37 .5) (30.7-35.7) (17.5-22.0)

6.0-6.4 42.8 ± 1.8 24.6 ± 1.9 32.9 ± 4.0 36.9 ± 3.4 21.4 ± 1.0 5 (40.0-45.1) (21.6-26.2) (30.7 -40.0) (31.2-40.0) (20.0-22.5) 41

Table 8: Morphometric proportions of Sebastes atrovirens larvae [x ± SD (range)]. See Table 6 for explanation.

SL SA/SL HL/SL SNL/HL ED/HL BD/SL n (mm)

4.0-4.4 38.1 ± 2.1 18.8 ± 2.1 31.0 ± 6.5 37.8 ± 6.1 14.1 ± 1.9 50 (34.0-43.9) (13.9-23.8) (12.5-44.4) (22.2-57.1) (11.3-18.6)

4.5-4.9 38.0± 2.7 19.4 ± 2.2 28.3 ± 5.4 38.1 ± 5.7 13.4 ± 2.1 38 (33.3-44.8) (15.2-28.4) (20.0-40.0) (25.0-50.0) (10.6-18.3)

5.0-5.4 39.9± 2.8 21.8 ± 3.1 27.6 ± 7.3 39.1 ± 2.2 14.3 ± 1.6 9 (35.8-45.0) (18.8-29.4) (16.6-40.0) (36.3-41.6) (11.3-16.6)

5.5-5.9 41.8 21.8 25.0 50.0 14.5 1

6.0-6.4 42.2 ± 1.8 24.7 ± 3.1 24.1 ±4.6 42.7 ± 7.4 17.5 ± 2.1 3 (40.9-44.2) (22.9-28.3) (21.4-29.4) (35.2-50.0) (16.3-19.9) 42

Table 9: Morphometric proportions of Sebastes rastrelliger larvae [x ± SD (range)]. See Table 6 for explanation.

SL SA/SL HL/SL SNL/HL ED/HL BD/SL n {mm)

4.0-4.4 34.2± 2.1 19.9 ± 1.7 30.0 ±4.5 37.2 ± 4.5 16.1 ± 1.3 5 (31. 8-36.5) (18.1-21.9) (25.0-33.3) (33.3-44.4) (14.6-18.1)

4.5-4.9 34.5 ± 2.1 18.9 ± 1.3 28.8 ±5.2 36.8 ± 5.4 15.6 ± 1.4 25 (29.7-37.5) (16.3-21.2) (22.2-37 .5) (30.0-55.5) (12.5-17.7)

5.0-5.4 35.3 ± 1.8 19.6±2.3 30.2 ±4.9 35.8 ± 5.2 14.7 ± 1.4 25 (31.4-38.0) (16.6-25.9) (20.0-36.3) (27 .2-44.4) (1 Ll-16.7)

5.5-5.9 38.0± 2.8 20.3 ± 1.9 32.8 ± 4.1 39.1 ± 4.8 15.9 ± 1.9 18 (32.7-41.0) (17.5-25.4) (25.0-41.6) (30. 7 -46.1) (12.0-18.6)

6.0-6.4 40.1 ± 1.7 24.4 ± 2.4 30.0 ± 4.1 35.3 ±4.8 17.4±1.9 15 (37.5-43.3) (20.9-28.3) (26.6-42.8) (26.6-42.8) (14.7-20.9)

6.5-6.9 40.2 ± 2.4 24.7 ± 1.3 30.8 ± 2.4 37.3 ± 3.7 17.9± 1.9 13 (34.8-43.0) (22. 7-26.4) (25.0-33.3) (31.2-43. 7) (15.4-21.4)

7.0-7.4 40.5 ±2.2 25.0 ± 1.7 30.9 ± 3.3 44.4 ± 5.0 20.3 ± 1.2 6 (37.1-43.0) (22.8-27.7) (25.0-33.3) (38.8-50.0) (18.5-22.2)

7.5-7.9 42.8 ± 1.4 26.3 ± 0.4 32.8 ± 6.3 39.3 ± 1.2 19.8 ± 2.2 3 (41.5-44.3) (25.9-26.6) (28.5-40.0) (38.0-40.0) (17.3-21.5) 43

Table 10: Morphometric proportions of Sebastes melanops larvae [x ± SD (range)]. See Table 6 for explanation.

SL SA/SL HLISL SNL/HL ED/HL BD/SL n (rnm)

3.5-3.9 37.7 ± 1.5 20.0 ± 1.1 35.9 ± 6.1 40.5 ± 4.1 16.3 ± 1.2 24 (35.1-40.5) (18.4-22.2) (25.0-42.8) (28.5-50.0) (12.8-19.4

4.0-4.4 37.6 ± 1.3 19.7 ± 1.4 37.0 ± 3.5 38.0± 2.9 15.6 ± 1.1 15 (35.0-40.0) (17 .5-22.5) (28.5-42.8) (33.3-42.8) (14.6-17.5) 44

Figure 1: Distribution of adult Sebastes mystinus, S. carnatus, S. atrovirens, S. rastrelliger, and S. melanops along the west coast of North America. 45

Alaska

Canada ...... s. mystinus

. · · · · S. carnatus • 1 • U.S.A . • ·---- S. atrovirens • Pacific • Ocean • S. rastrel/iger • • S. melanops •

Hawa11an" .• • ·• .Islands-.. () 46

Figure 2: Capture site of gravid Sebastes mystinus, and S. carnatus females. Rearing of all species (butS. melanops) occurred at the Monterey Bay Aquarium. 47

·.·

' .

. ~ . . . " '· . . : .. ' .. .. :: :.:' \ ..

·.' . ,• .. ·. . .

II. , . .. ; :Monterey Bay Aquarium

•' . ' '

.. : ., . ' Carmel , ... Bay

.. .. 48

Figure 3: Diagram of Sebastes larvae with axes used for morphometric measurements. SL= standard length; SA= snout-anus length; HL= head length; SnL= snout length; and ED= eye diameter. 49

_.J en 50

Figure 4: Diagram of Sebastes larva melanophore positions. Thirty three loci where melanophores were present are marked. Pectoral fin loci (7, 8, and 9) are shown separately. Locus: 1: symphysis of the lower jaw; 2: upper jaw; 3: lateral aspect of the lower jaw; 4: dorsal surface of the brain from the front to the back of the eye; 5: lateral surface of the brain from the front to the back of the eye; 6: dorsal surface from the back of the eye to the front of the cleithrum; 7: base of the pectoral fm; 8: blade of the pectoral [In; 9: margin of the pectoral fm; 10: ventral aspect of the gut; 11: dorsal aspect the gut; 12: posterio-dorsal surface of the gut; 13: posterio-ventral surface of the gut; 14: ventral membrane between anus and gut; 15: inside surface of the anus; 16: border of the anus; 17: outside surface of the anus; 18: dorsal midline from above the cleithrum to the the 3rd postanal myomere; 19: from the postanal myomere to the last myomere; 20: from the last myomere dorsally along the notochord to the hypural plates (if present); 21: first to 3rd or 4th postanal myomere; 22: third or 4th to 6th or 7th postanal myomere; 23: 7th to 19th postanal myomeres; 24: lateral aspect of the body from the 6th or 7th to the 19th postanal myomere; 25: from the last myomere ventrally along the notochord to the hypural plates (if present) or to the end of the notochord; 26: ventral aspect of the caudal fln; 27: lateral aspect of the body from the 1st to the 5th postanal myomere; 28: cleithral symphysis; 29: lower jaw symphysis; 30: fin fold behind the anus; 31: otic area; 32: olfactory lobe; 33: intemallocus above the dorsal surface the notochord from the 3rd to the 19th postanal myomere.

(Modified after Kendall and Lenarz, 1987) 20 4 31 6 1117 18 19 33 '

; . . ~. . -···..,7·,, .··:-~--·-- .. -~· .. .. ~··.····

1

25 26 10 12 13 14 16 15 30 21 27 22 23 24

8 52

Figure 5: Starved Sebastes mystinus larva. Note dorsoventrally shrunken head, "duck billed"jaw, and reduced body depth. 53 54

Figure 6 Growth curves of Sebastes mystinus (n=4), S. carnatus (n=3), S. atrovirens (n=2), S. rastrelliger (n=l), and S. melanops (n=l). n= number of females that gave birth. 55

0 tO lii "' "' iii"' g."' .§" ~ .:,,. ~ "' E g ~ -'ll" ~ "' l""' E" (Jj (Jj" (Jj"' (Jj (Jj 0 lO I ;' •I ' I t t t

--. (/) ;::... 10 0 "0 "\ ...... ·" \ "' ., (I) I 01 I <( I I I I

0 ' ' ' ' ' "' ' ' ' ' ' ' ~'"

0

(WW) lJlDUal pH!PUSlS 56

Figure 7: Growth series of Sebastes mystinus larvae. a.) 4.2 mm SL preflexion larva; b.) 4.8 mm SL preflexion larva; c.) 5.6 mm SL preflexion larva. 57

i1 ~~ ~\t ~~1\ ;._: .. (·· r· F: ~ r:~ :· {··... r<: f·· . .· ;···. ,. :-•. t:: ·:·. ['"· .... : f:3,.

~ . { : t"'(t •····••····... : 58

Figure 8: Melanophore densities (mean number per larva) at locus 4 and locus 6 of each size class for all species. 1= 3.0-3.4 mm; 2= 3.5- 3.9 mm; 3= 4.0-4.4 mm; 4= 4.5-4.9 mm; 5= 5.0-5.4 mm; 6= 5.5-5.9 mm; 7= 6.0-6.4 mm; 8= 6.5-6.9 mm; 9= 7.0-7.4 mm; 10= 7.5-7.9 mm. 59

II - Range Mean-- I~ Standard Deviation 30 Locus 4

13 6 3

"'~ 0 "'0 ~.r: 5 .!l" 0c. Ec z-:::J "' 10 71 ,,.III. :a"' 59 24 3 2i 6 5 10 I 3? 19 ljii 24 15 0 ~ i:I][I 1:~II I iii + +

1 2 3 4 5 2 3 4 5 6 7 3 4 5 3 4 5 6 7 8 9 10 2 3 S. mystinus S,camatus S.alrovirens S, rastrellif)ef S.me!anops

30 Locus 6 13 3

6 ;::"' 20 -0 0 ~.r: .J:l" 0Q 7 5 251 ,I, E c 8 1 z-:J .,"' 10 :a 59 24 10 II 32193 27II 6 5 4 3821 II 1 24 15 0 :lll:::m:~ III ~I i~II + +

i 2 3 4 5 2 j J 3 4 5 6 i d 9 ib 2 d S. mystinus S.camatus S.atrovirens S. rastreJJiger S.melanops

Species 60

Figure 9: Melanophore densities (mean number per larva) at locus 18 and locus 19 of each size class for all species. 1= 3.0-3.4 rnrn; 2= 3.5-3.9 rnrn; 3= 4.0-4.4 mrn; 4== 4.5-4.9 rnrn; 5= 5.0-5.4 rnrn; 6= 5.5-5.9 rnrn; 7= 6.0-6.4 rnrn; 8= 6.5-6.9 rnrn; 9= 7.0-7.4 rnrn; 10== 7.5-7.9 mm. 61

n - Range Mean- .. I- Standard Oev1aUon 30 Locus 18 13

20 15 6 3 7 10 18 p 3 21 1 3219 4 6 5 24 15 3+8 :&: iii I + + 0 + + + I,w:I :m:Il

1 2 3 4 5 2 3 4 5 6 7 3 4 5 3 4 5 6 7 8 9 10 2 3 S. mystim1s S.camatus S.atrovlrens S. rastrefiiger S.melanops

75 Locus 19

3 Ill 25 15 ~ 55 I 0 "0 ~.c: + -"" "-0 E c 35 z-" "' i!I1Ii :;;"' 1 6 1 3 59 24 1 0 7 5 32 3 15 + 5 4 38 21 5 I:m:IIIII Ii 24 15 0 + + ;;:s j[ X + +

2 3 4 5 2 3 4 5 6 7 3 4 5 3 4 5 6 7 8 9 10 2 3 S. mysHnus S.camatus S.a!rvvire(ls S. rastreffiger S.mefanops

Species 62

Figure 10: Growth series of Sebastes carnatus larvae. a) 3.5 mm SL yolksac larva; b) 4.3 mm SL preflexion larva; c) 6.0 mm SL preflexion larva. 63

.,

/\:"'"•,/r0 ./',v : ' ~ .. ~·.·.· '•, / 64

Figure 11: Growth series of Sebastes atrovirens larvae. a) 4.4 mm SL preflexion larva; b) 4.6 mm SL preflexion larva; c) 5.4 mm SL preflexion larva; d) 5.9 mm SL preflexion larva. 65

Sebastes atrovirens

2 Days Preflexion 4.4mm

10 Days Preflexion 4.6mm

19 Days Preflexion 5.4mm

~~·~·····";,~;:~'~,>~~·.~.-.··- '''"'' ..·:·:·.··-~.· '..·.:~·.·~.,·~.· . .,,.~,~.·.. :.~ ...···.·.' .. ---'7 ..·>·:. ~.\/.-~·.·..-~~.·.· ~.·.•• '-·"""·:,··.·'~A~.-'"'· .. ;;·'·7,·· .. ·/"~ .../-~·.·~ ~-~-., ----t·;i:}j}~.~~~?-::--,. -:::/~-·:;<<:;·. -. / - · . -.c,_.-...... ~._~~~-~-~,..:,_. ___ ··-¥~~~t~?E~~)T:~:::;;

- '·,_

28 Days Pre flexion 5.9mm 66

Figure 12: Growth series of Sebastes rastrelliger larvae. a) 4.6 mm SL yolksac larva; b) 5.8 mm SL preflexion larva; c) 6.4 mm SL flexion larva; d) 7.2 mm SL flexion larva; e) 7.0 mm flexion larva. 67

Sebastes rastrelliger

1 oay Preflexlon 4.6mm

20 Days Preflexlon s.amm

29 Days Flexion 6.4mm

34 Days Flexion 7.2mm 68

Figure 13: Growth series of Sebastes melanops larvae. a) 3.4 rum SL preflexion larva; b) 4.1 mm SL preflexion larva. 69

··~ .. ····~ ..-": ";., ·. : ·.. .· :··, ' '• ··.·: ..:: ('·, [·.:~~~ ?·.•

' ' ®'