NOAA Professional Paper NMFS 3

U.S. Department 

of Commerce

June 2004

Development of Kelp Rockfish, 

Sebastes atrovirens (Jordan and 

Gilbert 1880), and Brown Rockfish, 

S. auriculatus (Girard 1854), from 

Birth to Pelagic Juvenile Stage, with 

Notes on Early Larval Development 

of Black-and-yellow Rockfish, 

S. chrysomelas (Jordan and 

Gilbert 1880), Reared in the 

Laboratory (Pisces: Sebastidae)

William Watson

Larry L. Robertson
U.S. Department

of Commerce

Donald L. Evans

Secretary

National Oceanic

and Atmospheric

Administration

Vice Admiral

Conrad C. Lautenbacher Jr.,

USN (ret.)

Under Secretary for 

Oceans and Atmosphere

National Marine 

Fisheries Service

William T. Hogarth

Assistant Administrator 

for Fisheries

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Papers NMFS

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Editorial Committee

Dr. Andrew E. Dizon National Marine Fisheries Service

Dr. Linda L. Jones National Marine Fisheries Service

Dr. Richard D. Methot National Marine Fisheries Service

Dr. Theodore W. Pietsch University of Washington

Dr. Joseph E. Powers National Marine Fisheries Service

Dr. Tim D. Smith National Marine Fisheries Service

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NMFS series. 
NOAA Professional Paper NMFS 3

Development of Kelp Rockfish, 

Sebastes atrovirens (Jordan and 

Gilbert 1880), and Brown Rockfish, 

S. auriculatus (Girard 1854), from 

Birth to Pelagic Juvenile Stage, with 

Notes on Early Larval Development of 

Black-and-yellow Rockfish, S. chrysomelas

(Jordan and Gilbert 1880), Reared in the 

Laboratory (Pisces: Sebastidae)

William Watson

Larry L. Robertson

June 2004

U.S. Department of Commerce

Seattle, Washington
Suggested reference

Watson, William, and Larry L. Robertson. 2004. Development of kelp rockfish 

Sebastes atrovirens (Jordan and Gilbert 1880), and brown rockfish, S. auriculatus

(Girard 1854), from birth to pelagic juvenile stage, with notes on early larval development of black-and-yellow rockfish, S. chrysomelas (Jordan and Gilbert 1880), 

reared in the laboratory (Pisces: Sebastidae). NOAA Prof. Paper NMFS 3, 30 p.

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1

Introduction

Approximately 65 species of the viviparous rockfish genus Sebastes (Boehlert 

and Yoklavich, 1984; Moser, 1996b) 

occur in California waters (Eschmeyer 

et al., 1983; Eitner et al., 1999). Most 

are important sport and/or commercial fishery species (e.g., Lenarz, 1987; 

Lea et al., 1999; Aseltine-Neilson, 

2000). A live-fish fishery that targets 

nearshore species, especially rockfishes, developed in the 1980s and has 

grown greatly (e.g., Walters, 2001). 

Many of the rockfishes targeted by 

this fishery are members of the Sebastes

subgenus Pteropodus, which includes 

kelp (S. atrovirens), brown (S. auriculatus), gopher (S. carnatus), copper (S. 

caurinus), black-and-yellow (S. chrysomelas), calico (S. dalli), quillback 

(S. maliger), china (S. nebulosus), and 

grass (S. rastrelliger) rockfish (Hubbs 

and Schultz, 1933; Seeb, 1998). The 

effect of the fishery on stocks of the 

nearshore species is largely unknown, 

but it is not unreasonable to suppose 

that the depletion observed for other 

Sebastes species (e.g., Love et al., 1998; 

Ralston, 1998; Moser et al., 2000) may 

occur for these species as well. The 

Abstract—Larval kelp (Sebastes atrovirens), brown (S. auriculatus), and blackand-yellow (S. chrysomelas) rockfish were 

reared from known adults, to preflexion 

stage, nine days after birth for S. chrysomelas, to late postflexion stage for S. atrovirens, and to pelagic juvenile stage for S. 

auriculatus. Larval S. atrovirens and S. chrysomelas were about 4.6 mm body length 

(BL) and S. auriculatus about 5.2 mm BL 

at birth. Both S. atrovirens and S. auriculatus underwent notochord flexion at about 

6–9 mm BL. Sebastes atrovirens transform to 

the pelagic juvenile stage at about 14–16 

mm BL and S. auriculatus transformed at 

ca. 25 mm BL. Early larvae of all three 

species were characterized by melanistic 

pigment dorsally on the head, on the gut, 

on most of the ventral margin of the tail, 

and in a long series on the dorsal margin 

of the tail. Larval S. atrovirens and S. auriculatus developed a posterior bar on the tail 

during the flexion or postflexion stage. In 

S. atrovirens xanthic pigment resembled 

the melanistic pattern throughout larval 

development. Larval S. auriculatus lacked 

xanthophores except on the head until late preflexion stage, when a pattern 

much like the melanophore pattern gradually developed. Larval S. chrysomelas had 

extensive xanthic pigmentation dorsally, 

but none ventrally, in preflexion stage.

 All members of the Sebastes subgenus 

Pteropodus (S. atrovirens, S. auriculatus, 

S. carnatus, S. caurinus, S. chrysomelas, S. 

dalli, S. maliger, S. nebulosus, S. rastrelliger) 

are morphologically similar and all share 

the basic melanistic pigment pattern described here. Although the three species 

reared in this study can be distinguished 

on the basis of xanthic pigmentation, it 

seems unlikely that it will be possible to 

reliably identify field-collected larvae to 

species using traditional morphological 

and melanistic pigmentation characters.

Development of Kelp Rockfish, Sebastes atrovirens

(Jordan and Gilbert 1880), and Brown Rockfish, 

S. auriculatus (Girard 1854), from Birth to 

Pelagic Juvenile Stage, with Notes on 

Early Larval Development of Black-and-yellow 

Rockfish, S. chrysomelas (Jordan and Gilbert 1880), 

Reared in the Laboratory (Pisces: Sebastidae)

William Watson 

Larry L. Robertson

National Marine Fisheries Service, NOAA

Southwest Fisheries Science Center

8604 La Jolla Shores Drive

La Jolla, California 92037-1508

e-mail: William.Watson@noaa.gov

nearshore species recently were identified as requiring immediate fishery 

management attention, and they are 

included in the California Nearshore 

Fishery Management Plan of 2002 

(Walters, 2001).

Management of nearshore stocks 

requires information on population 

sizes; however, fishery-dependent data 

on population trends have not been 

reliable (e.g., McKee-Lewis, 1998; 

Walters, 2001). Ichthyoplankton can 

provide a fishery-independent measure of population trends (e.g., Moser 

and Watson, 1990; Moser et al., 2000), 

and has been demonstrated to be a 

useful tool for estimating population 

size (e.g., Lasker, 1985; Ralston et al., 

2003). For ichthyoplankton to be useful it must be possible to identify the 

larvae. Sebastes larvae are difficult to 

identify to species (e.g., Moser, 1996b) 

and currently no nearshore species 

is identified in ichthyoplankton samples. Published descriptions of complete larval development are available 

for only two of the Pteropodus species: 

S. dalli (Moser and Butler, 1981) and 

S. rastrelliger (Moreno, 1993; Laidig 

and Sakuma, 1998). Descriptions of 

partial developmental series are avail-
2

able for seven species: S. atrovirens (Moreno, 1993), 

S. auriculatus (DeLacy et al., 1964; Stahl-Johnson, 1985; 

Kendall1), S. carnatus (Moreno, 1993), S. caurinus

(DeLacy et al., 1964; Stahl-Johnson, 1985; Kendall1), 

S. chrysomelas (Wold2), S. maliger (DeLacy et al., 1964; 

Kendall1), and S. nebulosus (Kendall1). However, the 

existing literature provides no characters that allow 

identification of larvae to species through all developmental stages. The purpose of this paper is to provide 

additional information on development of S. auriculatus

and S. atrovirens from birth to pelagic juvenile stage, and 

early development of S. chrysomelas, based on laboratoryreared specimens. These are compared with larvae of 

the other Pteropodus species.

Materials and methods

Live adult S. atrovirens and S. chrysomelas were collected 

at San Miguel Island and S. auriculatus were collected 

near Point Conception, California, in March 1999. All 

fish were caught inshore (< 25 m depth) with hook and 

line or by SCUBA divers using dip nets. The fish were 

maintained aboard ship in two 500-l tanks with flowthrough (19 l/min) ambient seawater (12–15°C), and 

transported to the laboratory in those tanks with the 

water inlets and outlets closed. In the laboratory the fish 

were held in 1700-l tanks with sand-filtered, UV-treated, 

flow-through (19 l/min) chilled seawater (11±1°C). Obviously gravid females (three S. atrovirens, one S. auriculatus) were isolated in separate, flow-through (7.5 l/min, 

11±1°C) fiberglass tanks (360–680 l) that drained into 

fine-mesh (0.333 mm) collector baskets. All fish were 

fed to satiation twice weekly with thawed anchovy and 

squid, and formulated fish pellets (Bio-Oregon brood, 

12 mm). The isolated, gravid females showed no interest 

in feeding until after parturition.

An S. chrysomelas and an S. auriculatus extruded larvae 

aboard ship. The larvae were placed in 5-l beakers (~ 300 

larvae/l) maintained in water baths at 11.5°C (±0.5°C) 

with Lauda chillers (model RMS6). A small aquarium 

air pump provided light aeration and fluorescent lighting in the ship’s wet lab provided constant illumination. 

Larvae were fed copepod eggs and microplankton, but 

insufficient quantities were available and the larvae survived only 4–5 days after yolk exhaustion.

1 Kendall, A. W., Jr. 1989. Additions to knowledge of Sebastes larvae 

through recent rearing. U.S. Dep. Commer. NOAA, NMFS, NWAFC 

Processed Report 89-21, 46 p. Alaska Fisheries Science Center, 7600 

Sand Point Way NE, Seattle, WA 98115.

2 Wold, L. 1991. A practical approach to the description and identification of Sebastes larvae. Unpubl. M.Sc. Thesis, Calif. State Univ., 

Hayward, 88 p. California State University, Hayward, 25800 Carlos 

Bee Blvd., Hayward, CA 94542.

In the laboratory larvae were gently manually expressed from one of the gravid S. atrovirens and the 

gravid S. auriculatus. The other two gravid S. atrovirens

eventually released larvae naturally between noon 

and 3 PM for one, and sometime between 3:30 PM 

and 6:30 AM for the other. Manually expressed larvae 

were caught in 19-l buckets filled with 11°C seawater. 

Naturally spawned larvae were retained in the collector 

baskets and also were collected by partially draining the 

tanks and scooping larvae into 19-l buckets. All larval 

rearing was in static systems. Some larvae were stocked 

at ~ 200 larvae/l in black-plastic-wrapped, 5-l beakers in 

water baths cooled to 11°C with Lauda chillers. Others 

were stocked at ~ 50 larvae/l in various 19-l containers 

in a large water bath cooled to 11°C with an in-line titanium chiller (Aquanetics model AFC-4B). Fluorescent 

ceiling lights provided constant illumination, dimmed 

16 h/day when containers were partially covered. Light 

aeration was provided to each rearing container with a 

thin pipette attached to an aquarium air pump (Tetra 

model G-M). Detritus and moribund or dead larvae 

were siphoned from each container and ~ 5–20% of the 

water exchanged with fresh seawater at about 1–2 day 

intervals. Streptomycin and penicillin (Sigma Chemical 

Co.) were added at a dosage of 50mg/l each as a prophylactic treatment at about 10–12 day intervals.

From the day after parturition through notochord 

flexion, larvae were fed about daily with the marine 

rotifer Brachionus plicatilis, reared on algae (Tetraselmis

spp.). Wild plankton, predominantly copepod naupliar 

and copepodite stages and mollusc veligers, was added at 

irregular intervals, usually 2–3 times per week, depending on availability. Plankton was collected at the Scripps 

Institution of Oceanography pier using a 0.04 mm-mesh 

net. Plankton was filtered through a 0.15 mm-mesh sieve 

and the 0.04–0.15 mm fraction was fed to the larvae. Beginning late in the preflexion stage brine shrimp (Artemia salina) nauplii (Argent Labs) enriched with “HUFA” 

(Salt Creek, Inc.) were added to the diet, and the upper 

size limit of the wild plankton was increased to 0.3 mm. 

After notochord flexion, Artemia was the primary food, 

supplemented by wild plankton when available. Tetraselmis was added to the rearing containers occasionally to 

feed the prey organisms. Food densities varied according 

to availability, usually within the ranges of 5–12 Brachionus/ml, 5–10 Artemia/ml, and 4–6 plankters/ml. The 

19-l rearing containers were of three different colors 

(white, blue, black). Larvae in white containers did not 

feed successfully and none survived more than a few 

days after yolk absorption. Feeding was observed in the 

blue and black containers, but survival was poor in the 

black containers and only modest in the blue containers. 

Survival in the 5-l beakers also was modest.

For the descriptive series 1–100 larvae (usually 1–3) 

were collected at intervals of ~ 1–2 days during the first 
3

Figure 1

Locations of head spines referred to in the larval descriptions (modified 

from Ahlstrom and Moser, 1978). Abbreviations are: APO, anterior 

preopercular; IOP, interopercular; LIO, lower infraorbital; LOP, lower 

opercular; LPST, lower posttemporal; NA, nasal; NU, nuchal; PA, 

parietal; PPO, posterior preopercular; PRO, preocular; PSO, postocular; 

PT, pterotic; SC, supracleithral; SPO, supraocular; TM, tympanic; UOP, 

upper opercular; UPST, upper posttemporal.

two weeks and ~ 1–2 weeks thereafter. 

Specimens were preserved immediately 

in 2% sodium borate-buffered formalin. Totals of 96 S. atrovirens (4.4–14.6 

mm, preflexion–late postflexion stage), 

70 S. auriculatus (5.0–25.6 mm, preflexion–pelagic juvenile stage), and 110 

S. chrysomelas (4.4–5.4 mm, preflexion 

stage) were used for description. Ten 

field-collected S. atrovirens, obtained 

from the Scripps Institution of Oceanography Marine Vertebrates Collection 

(SIO H51-239:14.4–23.7 mm, late postflexion–pelagic juvenile stage), were examined to complete that series. Other 

species examined for comparison were 

20 S. caurinus (4.6–5.8 mm, preflexion 

stage) and 20 S. rastrelliger (4.8–6.4 mm, 

preflexion stage), reared at the Southwest Fisheries Science Center (SWFSC) 

experimental aquarium.

Most specimens were measured to 

the nearest 0.04 mm using a Wild M-5 

binocular microscope equipped with 

an ocular micrometer (a few moribund, 

shrunken larvae were used only for 

pigment descriptions). Dimensions 

measured were body length (BL), preanal length (PAL), head length (HL), 

head width (HW), snout length (SnL), 

eye diameter (ED), body depth (BD), 

pectoral (P1L) and pelvic (P2L) fin 

lengths, and the lengths of several head and pectoral 

girdle spines. These included: the two longest preopercular (PPO2, PPO3), upper and lower opercular 

(UOP, LOP), interopercular (IOP), parietal (PA), 

nuchal (NU), tympanic (TM), pterotic (PT), preocular 

(PRO), postorbital (PSO), first and second lower infraorbital (LIO1, LIO2), nasal (NA), upper and lower 

posttemporal (UPST, LPST), and supracleithral (SC) 

spines (Fig. 1). Spine terminology follows Moser and 

Ahlstrom (1978); dimensions are defined by Moser 

(1996a). Larval lengths always refer to BL of preserved 

larvae. Seven S. atrovirens, measured immediately after 

preservation and again 30–60 days later when all measurements for the descriptions were made, shrank an 

average 2% (range 0–5%) during the interval. Nine S. 

atrovirens (4.4–14.6 mm) and six S. auriculatus (4.8–25.6 

mm) were lightly stained with alizarin red S to aid in 

determining sequences of fin-ray and head spine formation. Occasionally, live or freshly preserved specimens 

(50 S. atrovirens, 62 S. auriculatus, 58 S. chrysomelas, 

mostly preflexion stage) were examined to document 

xanthophore patterns. Illustrations were made with the 

aid of a camera lucida. 

Description

Kelp rockfish (Sebastes atrovirens)

Morphology Larval S. atrovirens were 4.4–4.9 mm at 

birth (mean, mode = 4.6 mm) and began notochord 

flexion at 6.1–6.9 mm, 32–49 days later (11.4–12.0°C). 

Flexion was completed just after 8.6 mm, ~ 60 days after 

birth. The largest reared specimen, 14.6 mm, was late 

postflexion stage; among the field-collected specimens, 

one (14.4 mm) was transformation stage, one (15.5 

mm) was late postflexion stage, and the remainder 

(15.6–23.7 mm) were pelagic juveniles.

Larvae were moderately slender at birth, with a rounded head, short snout, short preanal length, and sac-like 

integument enclosing the trunk and anterior part of the 

tail (Table 1; Figs. 2A, 3A). A little yolk and/or single oil 

globule were present at birth; absorption was completed 

2–10 days later. The sub-dermal space gradually deflated 

during preflexion stage, and most body proportions 

gradually increased relative to BL throughout larval 

development (Table 1). Larvae had 25–26 myomeres 

(97% with 26): 6–8 preanal + 16–20 postanal (83% with 
4

A

B

C

D

Figure 2

Kelp rockfish, Sebastes atrovirens, lateral view. (A) 4.6 mm preflexion stage, day 1; (B) 4.6 mm preflexion stage, day 23; 

(C) 6.9 mm early flexion stage, day 52; (D) 8.0 mm mid-flexion stage, day 58; (E) 8.6 mm early postflexion stage, day 80; 

(F) 14.6 mm late postflexion stage, day 82; (G) 14.4 mm field-collected pelagic juvenile (SIO H51-239). Neuromasts are 

shown (with dotted lines) only on the preflexion-stage larvae.
5

E

F

G

Figure 2 (continued)

7 + 19) through flexion stage, shifting to 12 + 14 by late 

postflexion stage. 

Formation of the head spines began late in the preflexion stage (5.7 mm) with a preopercular spine (PPO3) 

at the angle of the posterior margin (Table 2). A much 

smaller spine (APO2) formed above and anterior to 

PPO3 at the end of preflexion stage. Simultaneously, or 

early in flexion stage (6.7–8.0 mm), a second posterior 

preopercular spine (PPO2) formed above PPO3. The 

third (lower) posterior spine (PPO4) formed by 7.1 mm 

and the second anterior spine (APO4) formed by 

8 mm. The last two preopercular spines, near its upper 

(PPO1) and lower (PPO5) ends, formed during postflexion stage by 11.9 and 14.6 mm, respectively. PPO3 

was largest until mid- to late postflexion stage (ca. 11.9–

14.6 mm), when PPO2 became nearly as long. In pelagic 

juveniles PPO2 was the longest spine (by ~ 20 mm), resulting from regression of PPO3. A small interopercular 

spine, which formed late in flexion stage, lacked a free 

distal end in some larvae and most pelagic juveniles. 
6

Table 1

Summary of measurements of kelp rockfish, Sebastes atrovirens, given as percentages of body length in 1 mm size classes. For 

each measurement mean values are given above, ranges below; n = number of specimens. Notochord flexion occurs within 

the 6.1–9.0 mm size classes; all but one of the specimens > 15.0 mm are pelagic juveniles (15.5 mm specimen = postflexion 

larva).

Size range n PAL BD HL HW SnL ED P1L P2L

 4.1–5.0 71 37 18 20 11 4 7 5 0

 36–43 14–20 18–23 10–14 3–7 7–8 3–7

 5.1–6.0 4 41 18 24 13 6 8 6 0

 40–43 17–19 22–26 11–16 5–8 7–9 5–7

 6.1–7.0 6 41 17 26 14 7 9 7 0.1

 40–43 11–20 24–28 13–15 5–9 8–10 6–7 0–0.3

 7.1–8.0 2 44 23 29 15 7 10 8 1

 43–45 21–24 27–30 14–17 7–7 9–10 8–8 1–1

 8.1–9.0 1 51 26 31 17 8 11 10 1

11.1–12.0 1 54 27 34 22 12 11 13 10

14.1–15.0 2 55 27 33 16 10 11 20 15

 55–56 26–27 31–34 15–18 9–11 10–11 18–22 13–16

15.1–16.0 3 57 27 35 15 12 11 21 17

 56–58 27–28 34–35 14–15 11–12 10–11 21–22 16–18

17.1–18.0 1 57 27 34 15 10 11 23 18

20.1–21.0 1 59 27 34 15 10 10 27 20

21.1–22.0 3 59 28 34 15 9 11 27 19

 58–59 28–29 33–35 15–15 9–10 10–11 26–28 19–20

23.1–24.0 1 59 29 33 15 10 10 26 19

Table 2

Summary of measurements of kelp rockfish, Sebastes atrovirens, head and pectoral girdle spines given as percentages of 

head length, in 1 mm size classes. For each measurement, mean values are given above, ranges below; n = number of 

specimens. Notochord flexion occurs within the 6.1–9.0 mm size classes; all but one of the specimens > 15.0 mm are 

pelagic juveniles (15.5 mm specimen = postflexion larva).

Size range (mm) n PPO2 PPO3 UOP LOP IOP PA NU PT PRO PSO LIO1 LIO2 NA LPST SC

 4.1–5.0 71 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

 5.1–6.0 4 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

 0–1

 6.1–7.0 6 0.3 5 0 0 < 0.1 1 0 1 0 0.2 0 0 0 0 0

 0–2 2–6 0–0.5 0–2 0–1 0–1

 7.1–8.0 2 4 10 0 0 0.2 5 0 2 0 2 0 0 0 0 0

 2–6 4–15 0–0.4 3–7 2–2 1–2

 8.1–9.0 1 7 25 0 0 0 10 0 3 0 2 0 0 0 0 0

11.1–12.0 1 15 20 0 0 0.4 12 1 2 0 2 0 0 0 4 2

14.1–15.0 2 13 15 1 0.4 0.4 6 3 1 0 0.4 1 1 1 3 3

 13–14 13–17 1–2 0–1 0.4–0.4 3–10 1–4 0–2 0–1 0–1 0–2 1–2 3–4 3–4

15.1–16.0 3 12 15 2 1 0.1 2 2 0 0 0 0.5 1 1 2 3

 12–13 14–15 2–3 1–2 0–0.4 0.4–4 1–3 0–1 0.4–1 1–1 2–3 2–4

17.1–18.0 1 12 10 3 2 1 0.3 1 0 0 0 1 1 2 1 2

20.1–21.0 1 10 5 4 3 0 0 2 0 0.3 1 1 1 2 1 3

21.1–22.0 3 12 8 3 2 0.1 0 2 0 0.3 1 0.4 0.5 2 1 2

 12–13 7–9 3–4 1–3 0–0.3 1–2 0–1 1–2 0–1 0.3–1 1–3 1–2 1–2

23.1–24.0 1 9 7 5 3 0 0 2 0 0.1 1 1 1 2 2 1
7

Figure 3

Kelp rockfish, Sebastes atrovirens, dorsal view. (A) 4.6 mm preflexion stage, day 1; (B) 4.6 mm preflexion stage, day 23; 

(C) 6.9 mm early flexion stage, day 52; (D) 8.0 mm mid-flexion stage, day 58; (E) 8.6 mm early postflexion stage, day 80; 

(F) 14.6 mm late postflexion stage, day 82. Neuromasts are shown only on the preflexion-stage larvae.

A

B

C

D

E

F
8

A small upper opercular spine formed late in postflexion stage, and transforming and pelagic juvenile specimens also had a smaller, lower opercular spine. 

A short, low pterotic ridge formed on the otic capsule during late preflexion or early flexion stage (5.6–

6.6 mm). A small pterotic spine formed on the ridge 

during flexion stage (6.1–7.1 mm) and persisted until 

late postflexion stage, but was absent in the pelagic 

juveniles. A pair of low, smooth parietal ridges formed 

during late preflexion or early flexion stage (5.7–ca. 6.9 

mm) and became finely serrate by 11.9 mm. A parietal 

spine formed posteriorly on each ridge during late preflexion to mid-flexion stage (5.9–7.1 mm) and a small 

nuchal spine formed just behind each parietal spine by 

11.9 mm. The parietal spine regressed during pelagic 

juvenile stage and by ~ 20 mm the nuchal spine was the 

only posterior spine on each parietal ridge. The postocular spine formed above the eye during notochord 

flexion at ca. 6.7–7.1 mm, and a smaller preorbital 

spine was present in pelagic juveniles ≥ ca. 21 mm. A 

lower infraorbital spine (LIO1) formed anteriorly on 

the lachrymal in some larvae ≥ 11.9 mm. Most pelagic 

juveniles had two lower infraorbital spines, but in some 

only the posterior spine (LIO2) was present. A pair of 

nasal spines formed at the end of the larval stage. Single 

supracleithral and posttemporal spines formed during 

postflexion stage, by 11.9 mm. 

The principal caudal-fin rays formed first, beginning 

late in preflexion stage at 5.0–5.5 mm. The full complement (8+7) was completed late in flexion stage (ca. 

8–8.5 mm), followed by formation of procurrent rays. 

These were added anteriorly from the last ray, and the 

full complement (10–11 + 9–11) was present by pelagic 

juvenile stage (Table 3). Pectoral-fin rays formed during late preflexion or early flexion stage, beginning 

at ~ 5–6 mm; addition of rays was ventrad and the full 

complement (16–18) was present in postflexion stage, 

Table 3

Fin-ray counts of reared kelp rockfish, Sebastes atrovirens. Abbreviations for fin rays are: D = dorsal, A = anal, P1 = pectoral, 

P2 = pelvic, CPRI = principal caudal, CPRO = procurrent caudal. Abbreviations for developmental stages are: Pr = preflexion, 

F = flexion, Po = postflexion.

BL (mm) Stage D A P1 P2 CPRI CPRO

 4.4 Pr (early) 0 0 0 0 0 0

 5.4 Pr (mid) 0 0 0 0 0 0

 5.0 Pr (late) 0 0 2 0 2 + 2 0

 6.4 F (early) 0 0 6 buds 5 + 5 0

 6.3 F (mid) 0 Anlage — buds 8 + 7 0

 8.4 F (late) anlage Anlage 8 buds 8 + 7 0 + 1

 8.8 Po (early) anlage 5 12 3 8 + 7 2 + 3

11.2 Po (mid) XIII,14 III,7 17 I,5 8 + 7 5 + 6

14.6 Po (late) XIII, 15 III,7 16 I,5 8 + 7 8 + 9

by 11.2 mm. Pelvic-fin buds formed during flexion 

stage at ca. 6.4–8.0 mm and rays formed in postflexion 

stage, beginning near 8.6 mm, with the full complement 

(I,5) present by 11.2 mm. Dorsal- and anal-fin anlagen 

formed during flexion stage, by ca. 8 mm, and segmented anal-fin rays formed late in the stage, by ca. 8.8 

mm. Anal-fin spines and dorsal spines and segmented 

rays formed during postflexion stage; full complements 

(D: XIII, 13–15; A: III, 6–7) were present in both fins 

by 11.2 mm. 

Pigmentation: Melanophores Larvae were moderately pigmented at birth (Figs. 2–4). Most lacked 

melanophores on the head and trunk (13% with a melanophore or two over the hindbrain, 18% with some 

on the trunk) but all had rows of melanophores dorsally 

and ventrally on the tail: usually 8–11 in a single dorsal 

row extending from myomere 9–17 through 21–23, and 

a broader ventral row, commonly of ~ 35–45 melanophores extending from myomere 7–8 through 22–24. 

Most larvae lacked melanophores laterally on the tail 

(pigment present in 20%, usually a melanophore ventrolaterally on one side between myomeres 21–23). The 

last 1–2 myomeres and notochord tip were unpigmented. The gut was heavily pigmented dorsally and there 

usually were 8–12 melanophores ventrally at mid-gut, 

with 1–2 on the hindgut adjacent to the anus.

Melanistic pigmentation gradually increased in all 

areas (Table 4). By day 2 or 3 melanophores were present above the myelencephalon and there usually were 

2–3 melanophores over the midbrain area, increasing 

after about day 14 to ≥ 12 covering the area by the end 

of preflexion stage (Fig. 3). There usually were two melanophores above the hindbrain through mid-flexion 

stage (ca. 7 mm), increasing in number to cover the 

myelencephalon by ca. 8 mm. Melanophores formed 

laterally on the cerebellum beginning 2–3 days after 
9

Figure 4

Kelp rockfish, Sebastes atrovirens, ventral view. (A) 4.6 mm preflexion stage, day 1; (B) 4.6 mm preflexion stage, day 23; 

(C) 8.0 mm mid-flexion stage, day 58; (D) 8.6 mm early postflexion stage, day 80; (E) 14.6 mm late postflexion stage, 

day 82. Neuromasts are shown only on the preflexion-stage larvae.

A

B

C

D

E
10

Table 4

Summary of melanophore distributions in larval kelp rockfish, Sebastes atrovirens. For number of melanophores, the range 

is given followed by the mode(s) in parentheses; for location of pigment, the area(s) where melanophores are located is 

given, followed by modal location(s) in parentheses (for trunk and tail pigment, location refers to the myomeres where 

melanophores are located). Modes are not given where clear modes were not apparent.

Larval stage

Pigment character Preflexion Flexion Postflexion

HEAD

 Forebrain area

 Number 0–3 (0) 0–3 (3) 6–12

 Location dorsal, anterior margin dorsal, anterior margin (dorsal) dorsal–upper 50%

 Midbrain area

 Number 0–31 (4) 16–many many

 Location dorsal, posterior margin dorsal, posterior margin dorsal, posterior margin–

(dorsal, posterior margin) (dorsal, posterior margin) upper 50%

 Hindbrain area

 Number 0–6 (2) several many

 Location dorsal, lateral, ventral (dorsal) dorsal, lateral, ventral dorsal, lateral, ventral 

(dorsal, lateral, ventral) (dorsal, lateral, ventral)

 Snout & upper jaw

 Number 0–1 (0) 0–2 (2) several

 Location anterior nostril, premaxilla, premaxilla, maxilla (premaxilla) anterior nostril, premaxilla, 

maxilla maxilla

 Lower jaw area

 Number 0–10 (0) 1–several few

 Location dentary, articular, retroarticular, dentary, articular, retroarticular dentary, articular, gular 

gular (dentary, retroarticular) (dentary, gular)

 Opercular area

 Number 0 0 several

 Location preopercle, opercle (opercle)

GUT AREA

 Dorsal

 Number many many many

 Location mid- & hindgut (mid- & hindgut) mid- & hindgut (mid- & hindgut) mid- & hindgut 

(mid- & hindgut)

 Lateral

 Number 0–several (0) few–many (many) many

 Location mid-gut area upper 25%–≥90% upper 50%–100%

 Ventral

 Number 7–21 (10, 14, 16) 9–many 5–many

 Location mid- & hindgut (mid- & hindgut) mid- & hindgut (mid- & hindgut) mid- & hindgut 

(mid- & hindgut)

 Isthmus

 Number 0–1 (0) 0–1 (0) 0

 Location near cleithra near cleithra

TRUNK & TAIL

 Dorsal, initial series

 Number 4–19 (10) 8–17 (13) 0–10

 Location 1–17 to 21–23 (13–23) 1–17 to 21–23 1–4 to 20–24

 Dorsal, secondary series

 Number 0 8–26 Many

 Location 1–5 to 18–20 1–4 to 24

 Ventral 

 Number ~ 30–72 (46, 48, 49) ~ 49–70 Many

 Location 6–9 to 22–24 (7–8 to 23) 7–8 to 23–24 (7–8 to 23–24) 8–17 to 24–25

 Lateral 

 Number 0–4 (0) 1–9 (4) 10– > 30

 Location 10–24, horizontal septum, 15–25, horizontal septum, 2–26 horizontal septum, 

ventrolateral (19–24) dorsolateral, ventrolateral dorsolateral, ventrolateral 

(24–25, horizontal septum) (22–26)

continued
11

Table 4 (continued)

Larval stage

Pigment character Preflexion Flexion Postflexion

 Internal

 Number 0–4 (1) Series Many

 Location 1–2 & 19–23, over &/or under 1–24, over & under notochord 1–24, over & under vertebral 

notochord (19–21, over (1–5 & 14–24, over notochord) column

notochord)

FINS

 Pectoral

 Number 0–10 1–several 0–several

 Location inner & outer surfaces of base, inner & outer surfaces of base, inner surface of base

proximally on membrane proximally on rays 

(inner surface of base) (inner surface of base, proximally 

on rays)

 Pelvic

 Number 0 0–2 (0) 0–7

 Location base base, rays I–2

 Dorsal

 Number 0 0 0–many

 Location between spines, soft-ray bases

 Anal

 Number 0 0–1 (0) 0–several

 Location last soft-ray base soft-ray bases (soft-ray bases)

 Caudal

 Number 0 0 9–several

 Location hypural margin, proximally 

on central rays

birth (present in 91%), gradually covering much of the 

lateral surface of the metencephalon and spreading posteriorly onto the myelencephalon by about mid-flexion. 

Basioccipital and/or pharyngobranchial melanophores 

usually formed late in preflexion stage (present in 16% 

≤ 2 weeks and in 100% ≥ 3 weeks). A few melanophores 

formed on the anterior margin and/or dorsally on the 

forebrain in mid-preflexion stage (ca. 5.4 mm). Subsequently, forebrain pigment changed little until postflexion stage, when more melanophores were added, 

covering the area by 14.6 mm. A melanophore formed 

near the mesial end of each premaxillary in 58% of late 

preflexion-stage specimens > 5.8 mm, commonly only 

on one side. One melanophore formed at the center of 

the upper jaw in 14% of specimens > 5.8 mm, and a few 

were present under the anterior end of each maxillary 

in all larvae > 7 mm. Some larvae ≥ 6.5 mm (22%) had 

≥ 1 melanophore at the anterior nostrils. On the lower 

jaw pigmentation ranging from a melanophore at the 

tip to a series along the anterior half usually formed after day 3 (present in 11% ≤ 7 days and in 59% of older 

larvae, including 86% of those ≥ 5 mm). Some larvae 

(12%) had 1–2 melanophores anteriorly on the gular area. Most larvae ≥ 5.8 mm (71%) had 1–3 melanophores 

on the articular near the retroarticular, and some larvae 

> ca. 6.5 mm (44%) had another 1–2 farther anteriorly 

on the ventral margin of the articular. The amount 

of articular pigment differed between left and right 

sides in 67% of the larvae. A melanophore formed on 

the isthmus near the cleithral symphysis in 20% of the 

larvae ≥ 5.7 mm. Pigmentation on the head increased 

during transformation–early juvenile stages, covering 

the upper 50–60% (sparsely laterally on the snout and 

opercle), extending over the jaws and onto the anterior 

gular area by 15.5–16 mm. The snout, jaws, and gular 

area were essentially fully pigmented by ~ 21 mm.

Melanophores spread ventrad from the dorsum of 

the gut, beginning at the anterior half of the mid-gut 

area, in mid–late preflexion stage (ca. 5.0–5.6 mm), 

and reached the ventrum in the anterior hindgut 

area by late preflexion to early flexion stage (ca. 5.4–

8.0 mm). By late flexion stage the gut was largely covered. Nearly all larvae had ≥ 1 melanophore on the peritoneum just anterior to the liver; in some preflexionand most later-stage larvae nearly the entire peritoneum 

was covered. The number of melanophores ventrally 

on the longitudinal midline of the gut increased a little 

during preflexion stage to 7–21 and by mid-flexion stage 

the ventral series usually was indistinguishable from the 

other gut pigment. 
12

The minimum number of melanophores dorsally on 

the trunk and tail increased from 4–5 early in preflexion stage to 9–10 late in the stage, but the maximum 

remained ~ 19 (Table 4). These usually were in a single 

row of 7–10 melanophores extending from myomere 

12–17 through 22–23. There commonly were 1–3 

melanophores on the trunk and/or tail isolated from 

the dorsal series during preflexion stage (present in 

68%), usually between myomeres 1–3 and 9–13: only 

on the trunk in 45%, only on the tail in 23%, and on 

both in 32% of the larvae having this pigment. During 

notochord flexion the dorsal melanophores became 

shallowly internal, beginning anteriorly, and by late 

postflexion stage (14.6 mm) none was visible externally. 

At about mid-flexion (ca. 7–8 mm) small melanophores 

formed between and adjacent to the original, larger 

dorsal melanophores on the tail, or both on the tail and 

at about mid-trunk. These small melanophores spread, 

extending from myomeres 1–4 through 20–24 during 

postflexion stage.

Most larvae lacked lateral melanophores on the trunk 

and tail during preflexion stage, but the proportion 

with some increased from ~ 17% early to ~ 36% late 

in the stage. When present, this pigment usually was a 

ventrolateral melanophore on one side in the vicinity 

of myomeres 19–24 until late in the stage when melanophores commonly formed on both sides, often with 

1–2 on the horizontal septum (present in 60%) in the 

same area. All flexion- and later-stage larvae had ≥ 1 

melanophore on the horizontal septum in the vicinity 

of myomeres 21–25. This pigment extended forward to 

mid-tail and expanded posteriorly during the pelagic 

juvenile stage but was obscured by other tail pigment 

after ~ 21 mm. Some flexion- and postflexion-stage 

larvae had melanophores dorsolaterally between myomeres 20–24 (25%) and ventrolaterally anywhere on 

the tail, commonly at myomeres 19–22 (38%). A bar 

formed here in transforming and early pelagic juvenile 

specimens. Late in postflexion stage melanophores 

spread ventrad from the dorsum, first in the vicinity of 

myomeres 7–9, then at myomeres 2–4. The anterior of 

these areas became a saddle extending from the nape 

to below the third dorsal-fin spine and ventrad nearly 

to the horizontal septum, by ~ 20 mm. The second became a saddle below D IV–VI or VII, extending to, or 

just below, the horizontal septum by ~ 20 mm. Two bars 

formed during transformation and early juvenile stages: 

below D IX–XI or XII, and below D 2 or 3–5 or 6; both 

neared the ventrum by ~ 20 mm. Thus, pelagic juveniles 

have five saddles and bars on the trunk and tail.

The number of ventral melanophores on the tail increased, with ~ 50–60 commonly present by the end of 

preflexion stage. These usually extended from myomere 

7–8 to 23–24 and usually were continuous with the pigment dorsally on the gut, although a small unpigmented 

gap sometimes separated the two areas (gap present in 

25%). The ventral melanophores became internal, beginning anteriorly, during postflexion stage, with those 

on the caudal peduncle remaining at least partially 

external to at least early pelagic juvenile stage.

Melanophores formed internally above the notochord late in preflexion stage (by ca. 6 mm), beginning 

with 1–2 in the vicinity of myomeres 19–21 and often 

with another at myomere 1 or 2 (present in 60%), and 

spread from both sites to extend the full length of the 

vertebral column by the end of flexion stage. Melanophores formed under the notochord/vertebral column 

at about myomere 23 by late preflexion stage in some 

larvae, but were absent until postflexion stage in others (present in 45% ≥ 6 mm). These melanophores 

remained at myomeres 21–24 in some larvae, but extended the full length of tail in others, beginning at 

mid-flexion to postflexion stage.

One or more melanophores formed proximally on 

the mesial surface of one or both pectoral-fin bases 

late in preflexion stage (ca. 5.7 mm). During flexion 

stage melanophores covered the mesial surface in 

some larvae, but in others the fin bases were sparsely 

pigmented or even unpigmented (unpigmented in 

10% of flexion- and postflexion-stage specimens). One 

to a few melanophores formed on the lateral surface 

of the pectoral-fin base in 23% of larvae ≥ 6 mm. A few 

melanophores formed proximally on some upper or 

lower pectoral-fin rays, or on both areas, in half of the 

late preflexion-stage and older larvae ≥ 6 mm; however, 

pectoral fin-ray pigmentation was not consistently present until pelagic juvenile stage, by ca. 20–21 mm. A little 

pigment was present on each pelvic-fin base in 30% 

of the late flexion- and postflexion-stage larvae, and a 

postflexion-stage specimen had a few melanophores 

proximally on the pelvic-fin rays (Table 4).

Melanophores first formed on the dorsal fin between 

dorsal spines II–V and at the bases of the segmented 

rays late in postflexion stage. Pigmentation increased 

rapidly, forming a dense bar at D I–III and sparser bars 

at D IV–V through VI–VII, D IX–XI or XII, and D 2–3 

through 5–6, by early pelagic juvenile stage (< 16 mm). 

Fin bars first appeared just before the corresponding 

saddles and bars on the body. A distal band connected 

the second and third bars on the spinous dorsal, and the 

fourth bar extended posteriorly to about D 9, in pelagic 

juvenile stage by ca. 21 mm. All but about the distal 

10–20% of the segmented-ray portion of the dorsal fin 

was pigmented by 23.7 mm. Melanophores formed at 

the bases of all, or all but the last segmented anal-fin 

rays, late in postflexion stage (≥ 14.6 mm). A band first 

appeared on the anal fin during transformation, tapering posteriorly from most of the length of A I down to 

the base of the last ray by early pelagic juvenile stage 

(< 16 mm). Melanophores formed on the hypural mar-
13

gin and, in one specimen, proximally on some central 

rays by 11.9 mm. Caudal fin-ray pigmentation in transformation stage was largely proximal on the principal 

rays at first, but extended nearly the full length of all 

principal rays and most procurrent rays by ca. 16 mm.

Pigmentation: Other chromatophores At birth 

xanthophores (yellow) were distributed much like the 

melanophores, but were more numerous than melanophores on the dorsum and less so on the ventrum. Xanthophores (2–4) covered much of the dorsal surface 

of the midbrain area and continued posteriorly as two 

rows to myomere 21–23, except that commonly there 

were none above the hindbrain. Xanthophores flanked 

the melanophores dorsally on the tail, expanding and 

spreading downward posteriorly. Ventrally on the tail, 

they formed a sparse row interspersed along the full 

length of the melanophore series in some larvae, but 

more commonly were posterior only, often expanded 

and spreading upward posteriorly. The expanded dorsal and ventral xanthophores commonly formed a bar 

at about myomeres 22–23 (present in 75%). On the 

gut, xanthophores were limited to the dorsum at midgut and anteriorly on the hindgut. The oil globule was 

golden yellow.

By late preflexion stage (ca. 5 mm) yellow pigment 

covered much of the midbrain area and was present 

internally in the otic capsule and anterolaterally on 

the hindbrain, but was absent externally over the hindbrain area in most larvae. By late flexion stage (ca. 8.8 

mm) it covered the mid- and hindbrain, and extended 

anteriorly over the forebrain and posteriorly over the 

spinal chord to the antepenultimate vertebra. The entire dorsal and dorsolateral surfaces of the head were 

covered by late postflexion stage (15.2 mm). Series of 

small, orange xanthophores formed on the dentaries 

and articulars during preflexion stage, and during 

postflexion stage more formed at the bases and distally along the margins of the central two preopercular 

spines. A few (yellow) were present on the anterior 

preopercular margin of one flexion-stage specimen. By 

late postflexion stage dorsal xanthophores spread onto 

the snout, forming clusters on the premaxillary ascending processes and spreading along the proximal half of 

each maxilla. In the latter part of postflexion stage a 

few xanthophores (orange) formed anteriorly on the 

gular area, a few (yellow) formed at the bases of the supracleithral and posttemporal spines, and guanophores 

formed on the opercular area. By late postflexion (15.2 

mm) the entire opercular area (except the preopercle) 

was bright silver.

External xanthic pigmentation on the trunk and tail 

changed little in preflexion stage. During notochord 

flexion xanthophores spread ventrolaterally from the 

dorsum at the first 1–3 myomeres and on the tail, extending progressively farther down the sides beginning 

at myomeres 14–18 and reaching the horizontal septum 

at about myomere 19–20. Xanthophores also proliferated ventrally on the tail during flexion stage, extending 

anteriorly and spreading dorsolaterally, reaching progressively farther up the sides posteriorly and meeting 

the dorsolateral pigment at about myomere 19, thus expanding the tail bar to about myomeres 19–24. The bar 

was predominantly yellow, with some orange pigment. 

During postflexion stage a xanthophore series formed 

on the horizontal septum of the tail, the bar spread 

anteriorly another 2–3 myomeres, and xanthophores 

covered (sparsely) the trunk to about myomere 5–6. A 

few xanthophores (yellow and orange) were present on 

the central hypural area in a flexion-stage specimen.

Xanthophores covered the gut during flexion and 

postflexion stages. Guanophores formed on the abdominal area during the latter part of postflexion stage; 

by late postflexion (15.2 mm) the entire area was bright 

silver. There were no xanthophores on the finfolds during preflexion stage; later a few formed on the pectoral 

fins near the bases of some rays in some larvae. A few 

xanthophores were present on the preanal finfold of a 

flexion-stage specimen.

Brown rockfish (Sebastes auriculatus)

Morphology Larval S. auriculatus were 5.0–5.3 mm 

long (mean 5.2 mm) at birth and completed yolk 

absorption in 9–13 days (5.4–5.8 mm). Notochord 

flexion began at 6–8 mm (32–45 days after birth) and 

was completed before 10 mm. Transformation to the 

pelagic juvenile stage was at ca. 25 mm, ~ 90 days after 

birth (11.5°C).

The S. auriculatus larvae were larger at birth and 

transformation but otherwise morphologically similar 

to S. atrovirens (Figs. 5–7; Tables 5–8). Larval S. auriculatus had 26–27 myomeres (92% with 26): 7–8 preanal 

+ 18–20 postanal during preflexion stage (90% with 

7 + 19), shifting to 10 + 16 in postflexion stage. Head 

and pectoral girdle spine development was similar in S. 

auriculatus and S. atrovirens, except that interopercular 

spines formed later, during postflexion stage, in S. auriculatus (Table 6), and pelagic juvenile S. auriculatus

had tympanic spines (lacking in S. atrovirens), but lacked 

preorbital spines (present in S. atrovirens).

Principal caudal- and pectoral-fin rays began to form 

early in flexion stage (~ 7 mm) and full complements 

(8+7 and 15–19, respectively) were present by early 

postflexion stage (Table 7). Procurrent caudal-fin rays 

formed in postflexion stage; addition was anteriorly 

from the posterior-most rays and the full complement 

(9–11 + 9–12) was present by pelagic juvenile stage. 

Pelvic-fin buds formed early in flexion stage, and dorsaland anal-fin anlagen in mid-flexion (Table 7), with all 
14

A

B

C

Figure 5

Brown rockfish, Sebastes auriculatus, lateral view. (A) 5.2 mm preflexion stage, day 1; (B) 5.8 mm preflexion stage, day 5; 

(C) 8.2 mm early flexion stage, day 45; (D) 10.6 mm postflexion stage, day 59; (E) 25.6 mm, pelagic juvenile, day 94. Neuromasts are shown only on the preflexion-stage larvae.

fin-rays (I,5; XIII,12–15; III,5–8, respectively) present 

early in postflexion stage. 

Pigmentation: Melanophores Larval S. auriculatus

were moderately pigmented at birth (Figs. 5–7), most 

commonly with about seven melanophores dorsally on 

the head, 9–16 dorsally on the tail between myomeres 

9–11 and 22–23, and a single row of 27–31 ventrally 

on the tail from myomere 9 through 23. The last 1–2 

myomeres and notochord tip were unpigmented. At 

birth 45% had pigment on one or both sides of the tail, 

usually consisting of a melanophore between myomeres 

18–21. All had heavy pigment dorsally on the gut, and 

about five melanophores ventrally at mid-gut plus one 

on the hindgut.

Pigmentation increased on all areas (Table 8). The 

upper ~ 25–50% of the midbrain area was covered by 

early flexion stage (Figs. 5, 6). Pigmentation over the 

myelencephalon increased from the initial 0–4 (usually 

2–3) melanophores to nearly cover it by flexion stage, 

and melanophores formed laterally on the cerebellum 

by day 2–3 and covered much of the metencephalon by 

early flexion stage. Melanophores formed at the anterior margin of the forebrain as early as day 4 in some 
15

Figure 5 (continued)

D

E

Table 5

Summary of measurements of brown rockfish, Sebastes auriculatus, given as percentages of body length in 1 mm size classes. 

For each measurement mean values are given above, ranges below; n = number of specimens. Notochord flexion occurs 

within the 6.1–9.0 mm size classes; the 25.6 mm specimen is a pelagic juvenile.

Size range n PAL BD HL HW SnL ED P1L P2L

 4.1–5.0 2 41 26 24 14 5 9 5 0

 39–42 26–26 23–24 14–14 5–5 9–9 4–6 0–0

 5.1–6.0 56 40 23 23 13 6 9 5 < 0.1

 38–48 16–26 22–32 12–17 5–9 8–12 4–9 0–0.4

 6.1–7.0 1 42 19 27 14 8 10 8 0

 7.1–8.0 2 47 22 30 15 9 10 8 1

 46–48 20–23 29–31 15–16 9–9 10–11 8–8 1–1

 8.1–9.0 1 44 20 28 13 9 9 8 0.4

10.1–11.0 1 55 27 35 21 11 12 8 6

25.1–26.0 1 62 28 34 20 10 9 24 18

larvae, but not until flexion stage in most. Forebrain 

melanophores were primarily dorsal during flexion 

stage, covering much of the upper half by late postflexion stage. Melanophores formed on the basioccipital by 

day 2, usually posteriorly on its dorsal and/or ventral 

margins. In ~ 50% of preflexion-stage larvae ≥ 4 days old 

this pigment spread forward and/or posteriorly under 

the anterior end of the notochord to the level of future 

vertebrae 1–3.

After the first week 58% of preflexion-stage larvae, 

and all later-stage specimens, had melanophores anteriorly near the bases of the premaxillary ascending 
16

Figure 6

Brown rockfish, Sebastes auriculatus, dorsal view. (A) 5.2 mm preflexion stage, day 1; (B) 5.8 mm preflexion stage, day 5; 

(C) 8.2 mm early flexion stage, day 45; (D) 10.6 mm postflexion stage, day 59. Neuromasts are shown only on the preflexion-stage larvae.

A

B

C

D
17

Figure 7

Brown rockfish, Sebastes auriculatus, ventral view. (A) 5.2 mm preflexion stage, day 1; (B) 5.8 mm preflexion stage, day 5; 

(C) 8.2 mm early flexion stage, day 45; (D) 10.6 mm postflexion stage, day 59. Neuromasts are shown only on the preflexion-stage larvae.

A

B

C

D
18

Table 6

Summary of measurements of brown rockfish, Sebastes auriculatus, head and pectoral girdle spines given as percentages 

of head length, in 1 mm size classes. For each measurement mean values are followed by ranges in parentheses. Number 

of specimens is given in parentheses for each size class. Notochord flexion occurs within the 6.1–9.0 mm size classes; the 

25.6 mm specimen is a pelagic juvenile.

Size range

Spine 4.1–5.0 (2) 5.1–6.0 (56) 6.1–7.0 (1) 7.1–8.0 (2) 8.1–9.0 (1) 10.1–11.0 (1) 25.1–26.0 (1)

PPO2 0 < 0.1 (0–2) 2 6 (5–7) 7 13 11

PPO3 0 0.1 (0–5) 8 11 (10–12) 11 18 12

UOP 0 0 0 0 0 0 4

LOP 0 0 0 0 0 0 1

IOP 0 0 0 0 0 1 0

PA 0 0.1 (0–4) 3 6 (6–7) 5 11 0

NU 0 0 0 0 0 2 3

PSO 0 < 0.1 (0–0.5) 1 1 (1–1) 1 2 2

PT 0 < 0.1 (0–2) 3 4 (3–5) 4 4 1

TM 0 0 0 0 0 0 2

LIO1 0 0 0 0 0 1 1

LIO2 0 0 0 0 0 0.5 0.5

NA 0 0 0 0 0 0 2

UPST 0 0 0 0 0 0 2

LPST 0 0 0 0 0 3 1

SC 0 0 0 0 0 0 3

Table 7

Fin-ray counts of reared brown rockfish, Sebastes auriculatus. Abbreviations for fin rays are: D = dorsal, A = anal, P1 = 

pectoral, P2 = pelvic, CPRI = principal caudal, CPRO = procurrent caudal. Abbreviations for developmental stages are: Pr = 

preflexion, F = flexion, Po = postflexion, PJ = pelagic juvenile.

BL (mm) Stage D A P1 P2 CPRI CPRO

 4.8 Pr 0 0 0 0 0 0

 7.0 F (early) 0 0 3 buds 5 + 5 0

 7.2 F (mid) anlage anlage 6 buds 5 + 5 0

 6.7 F (late) anlage anlage 14 buds 6 + 6 0

10.6 Po XII,14 III,7 18 I,5 8 + 7 5 + 5

25.6 PJ XIII,13 III,7 18 I,5 8 + 7 9 + 9

processes, and on the ascending processes in ~ 33% of 

the flexion-stage and older larvae. A melanophore was 

present on each maxilla in a preflexion-stage specimen 

(13 days), and most (86%) flexion-stage and later larvae 

had some pigment anteriorly under the maxillae. The 

snout usually was unpigmented into flexion stage (two 

flexion-stage larvae had a melanophore at the anterior 

margin of the nasal capsule), but by mid-postflexion 

stage melanophores nearly covered it to the level of 

mid-eye (Fig. 6). Melanophores formed on the lower 

jaw between days 3–7, usually anteriorly and laterally, 

but often not in both locations. About 75% of preflexion-stage larvae ≥ 5 days old had 1–2 melanophores on 

the lower margin of the articular, and later-stage larvae 

commonly had them along its full length. A melanophore was present on the retroarticular, usually on its 

inner surface, in 43% of preflexion larvae ≥ 11 days old 

and in 89% of later-stage larvae. A late preflexion-stage 

specimen had a melanophore anteriorly on the gular 

membrane, and one each of flexion and postflexion 

stage had several. About 75% of preflexion-stage larvae 

≥ 7 days old had 1–2 melanophores on the isthmus near 

the cleithral symphysis, and all later-stage larvae typically 

had 2–3. Larvae ≥ 7 days old usually had a melanophore 
19

Table 8

Summary of melanophore distributions in larval brown rockfish, Sebastes auriculatus. For number of melanophores, the 

range is given followed by the mode(s) in parentheses; for location of pigment, the area(s) where melanophores are 

located is given, followed by modal location(s) in parentheses (for trunk and tail pigment, location refers to the myomeres 

where melanophores are located). Modes are not given where clear modes were not apparent.

Larval stage

Pigment character Preflexion Flexion Postflexion

HEAD

Forebrain area

 Number 0–1 (0) 2–several several

 Location anterior or lateral dorsal–upper 50% (dorsal) upper 50%

Midbrain area

 Number 1–11 (3–4) several several

 Location dorsal, lateral (dorsal) upper 25–50% upper 50%

Hindbrain area

 Number 0–6 (3–4) several several

 Location dorsal, lateral (dorsal) upper 25–50% upper 50%

Snout & upper jaw

 Number 0–3 (0, 1) 0–4 (0, 2) 2–many

 Location premaxilla, maxilla (premaxilla) nasal capsule, premaxilla, anterior nostril–upper 50%, 

maxilla (premaxilla, maxilla) premaxilla, maxilla

Lower jaw area

 Number 0–8 5–several many

 Location dentary, articular, retroarticular, dentary, articular, retroarticular, dentary, articular, retroarticular, 

gular (dentary) gular (dentary, articular, gular (dentary, articular, 

retroarticular) retroarticular)

Opercular area

 Number 0–1 (0) 3–many many

 Location preopercle preopercle, opercle preopercle, opercle

GUT AREA

Dorsal

 Number several many many

 Location full length full length full length

Lateral

 Number 0–several several many

 Location mid-gut/hindgut area entire area entire area

Ventral

 Number 3–23 (indistinguishable from general (indistinguishable from general 

gut pigmentation) gut pigmentation)

 Location mid- & hindgut

Isthmus

 Number 0–2 1–4 2–3

 Location near cleithra near cleithra near cleithra

TRUNK & TAIL

Dorsal, initial series

 Number 0–20 (16) 11–15 (12, 14) 10–13

 Location 1–17 to 22–24 (1 & 11–22) 10–13 to 22–25 (12 to 22–25) 1–11 to 23–24

Dorsal, secondary series

 Number 0 several many

 Location 1–23 1–24

Ventral

 Number 16–35 (28) 28–40 ≥ 44

 Location 7–9 to 22–25 (9–23) 7–8 to 23–24 (8–23) 10–25

Lateral

 Number 0–4 (1) 2–9 (9) 3–many

 Location 12–23 (20) 1–3 & 8–24 (1 & 10 to 20–21) 1–26 (13–24)

Internal 

 Number 0–2 (0) series series

 Location 18–24, over or under notochord, 1–24, over & under notochord/ 1–25, over, under & lateral to 

or lateral (20–22, over notochord) vertebral column vertebral column

continued
20

Table 8 (continued)

Larval stage

Pigment character Preflexion Flexion Postflexion

FINS

Pectoral

 Number 0 0–several 10–many

 Location inner & outer surfaces of base, inner & outer surface of base, 

proximally on blade or rays on rays

(inner & outer surfaces of base) 

Pelvic

 Number 0 0–2 (0) 0–several

 Location base base, rays

Dorsal

 Number 0 0 0–many

 Location soft-ray bases

Anal

 Number 0 0–5 (0) 0–several

 Location finfold near origin of anlage soft-ray bases

Caudal

 Number 0–1 (0) 0–1 (0) 0–several

 Location finfold near hypurals proximally on a central ray proximally on principal rays

on the lower (anterior) part of the preopercle, and 55% 

of flexion-stage and later specimens had one near the 

preopercular angle and/or on the upper part of the 

preopercle. Most (78%) flexion- and postflexion-stage 

larvae had melanophore(s) at the base(s) of ≥ 1 posterior preopercular spine(s), commonly at PPO3. The 

opercular melanophore patch characteristic of pelagic 

juveniles began to form during postflexion stage. By 

pelagic juvenile stage the upper half of the head was 

nearly fully pigmented, most densely on its upper quarter, with a light peppering over the jaws and gular area, 

and a more or less prominent patch near the upper end 

of the opercle (Fig. 5E).

A melanophore or two formed on the peritoneum 

just anterior to the yolk and/or oil globule during the 

first day in most larvae, and spread ventrad to become 

continuous with the ventral pigmentation on the gut by 

late preflexion stage (Fig. 5). Melanophores began to 

spread ventrad from the dorsum of the gut (especially 

in the posterior mid-gut/anterior hindgut vicinity) on 

about day 3, and nearly reached the ventrum by late 

preflexion stage. By late flexion stage most of the gut 

was pigmented, more sparsely ventrolaterally. Beginning 

about day 7, some melanophores in the vicinity of the gas 

bladder extended up to the notochord. The number of 

melanophores ventrally on the gut gradually increased 

during preflexion stage, forming a broad longitudinal 

row by day 4 and becoming indistinguishable from the 

other gut pigmentation by late flexion stage (Fig. 7).

The location and number of dorsal melanophores on 

the tail changed little during preflexion stage (Figs. 5, 

6): larvae commonly had 12–16 melanophores between 

myomeres 11–22 (Table 8), mostly in a single row, except the row commonly was doubled posteriorly at one 

or two myomeres. The dorsal melanophores typically 

were more widely spaced anteriorly. Some preflexionstage larvae ≥ 7 days old (58%) also had some dorsal 

pigment on the trunk, usually a melanophore at the first 

myomere. Early in flexion stage an irregularly spaced 

dorsolateral row of smaller melanophores formed on 

each side of the initial series, usually originating simultaneously in the vicinities of myomeres 1–3 and 7–10. 

By mid-flexion stage the dorsolateral melanophores 

spread posteriorly on the trunk and formed in the vicinity of myomeres 14–23, and by late flexion they filled in 

between sites and merged into the original marginal 

series posteriorly. During postflexion stage the original marginal series became shallowly internal and the 

number of dorsolateral melanophores increased, forming a 2–3 melanophore-wide band on each side of the 

dorsal margin from the nape to mid-caudal peduncle 

(Fig. 6D). By 10.6 mm this pigmentation was sparse at 

the first 1–2 myomeres but dense along the bases of the 

segmented dorsal-fin rays.

The proportion of larvae with lateral melanophores 

on the tail rapidly increased after birth to 100% within 

two weeks. The number of lateral melanophores increased to 2–9 on each side by flexion stage (Table 8), 

then changed little through mid-postflexion stage 

(Fig. 5D). These melanophores were dorso- and ventrolateral, and on the horizontal septum, usually between 

myomeres 10–21, with most between myomeres 18–21. 
21

About 10% of preflexion-stage larvae ≥ 7 days old and 

just over half of flexion-stage larvae also had 1–2 melanophores dorsolaterally or on the horizontal septum 

in the vicinity of myomeres 1–3. In the latter part of 

postflexion stage melanophores spread forward on the 

horizontal septum, extending along its full length by 

pelagic juvenile stage (Fig. 5E). Others spread from the 

dorsum onto the sides of the trunk and tail, combining 

with the lateral pigment to cover much of the upper 

half of the body (densest on the upper quarter), and 

formed five saddles and bars in early pelagic juvenile 

stage (Fig. 5E). The first saddle extended diagonally 

from below D I–III to the posttemporal area, the second 

from below D IV–VIII to near the horizontal septum, the 

third from below D IX–XII to just below the horizontal 

septum, the fourth (a bar) from below D 2–11 nearly to 

the ventrum, and the last bar covered much of the caudal peduncle. Pigmentation in the saddles and bars was 

densest dorsally and increasingly sparse ventrad.

The number of ventral melanophores on the tail 

increased with development (Table 8). These were 

mostly in a single row, although the proportion of larvae 

with the row doubled at some myomeres posteriorly 

increased to just over 40% by flexion stage. Melanophores in the ventral row usually were small anteriorly, progressively more expanded posteriorly to about 

myomere 20–22, then progressively more contracted 

again (Fig. 7). Early in postflexion stage smaller melanophores formed, interspersed with the original series, 

and by mid-postflexion stage the original melanophores 

were shallowly internal except on the caudal peduncle. 

The smaller melanophores formed a 1–2 melanophorewide row on each side of the anal-fin base in the latter 

part of postflexion stage (Fig. 7D).

Larvae rarely had internal pigment in the trunk or 

tail during the first 2–3 days, but 58% of preflexionstage larvae ≥ 7 days old had 1–2 melanophores, usually 

above the notochord at about myomeres 20–22. During 

flexion stage a series of melanophores formed above 

the spinal cord, initially in the vicinities of myomeres 

1–2, 15–17, and 21–24. Two or three melanophores 

formed under the notochord at myomeres 19–21 early 

in flexion stage, and spread forward to myomeres 14–18 

and posteriorly to myomere 23 by late in the stage. 

Others spread forward dorsally along the notochord to 

myomeres 14–16, and some formed over the first 1–4 

vertebrae by late flexion stage. A few formed elsewhere 

internally, primarily dorsally and ventrally in the vicinity 

of myomeres 18–23, during postflexion stage.

A preflexion-stage larva had a ventral melanophore in 

the caudal finfold (Fig. 5B); others lacked fin pigmentation before flexion stage (Table 8). During flexion 

stage melanophores formed on the inner surface of 

the pectoral-fin base, usually proximally on its lower 

part, and some usually formed on the lateral surface. 

A few formed on the pectoral-fin blade, usually on its 

upper half, during early- to mid-flexion stage. One or 

two melanophores formed on each pelvic-fin base in 

some flexion- and postflexion-stage larvae (44%) and 

beginning about mid-postflexion stage, up to several 

formed on the membranes between the pelvic-fin rays. 

Melanophores formed on the bases of the segmented 

dorsal-fin rays at about mid-postflexion stage, but none 

formed on the fin rays or membranes before late postflexion or transformation stage. In the 25.6 mm pelagic 

juvenile all but about the distal 20–40% of the spinous 

dorsal fin and proximal 25% of the segmented ray portion of the fin were pigmented (except none on the last 

ray). A flexion-stage specimen had a few melanophores 

anteriorly on the anal finfold, but otherwise the anal fin 

was unpigmented before mid-postflexion stage, when 

melanophores formed at the bases of the segmented 

rays. The pelagic juvenile had a few melanophores at 

A III–1, and one each proximally on some other segmented rays. One or two melanophores were present on 

the lower hypural margin in some flexion-stage larvae, 

and by mid-postflexion stage there were several on the 

hypural margin. By pelagic juvenile stage a narrow bar 

extended along the hypural margin, and there were 

several melanophores proximally along each principal 

caudal-fin ray and a few farther distally on the fin.

Pigmentation: Other chromatophores Xanthophore 

and melanophore patterns differed from one another 

through mid-preflexion stage, but became more alike 

in older larvae. At birth, most larvae had a few yellow 

xanthophores dorsally over the midbrain, occasionally 

1–2 over the hindbrain, and none elsewhere. The oil 

globule was pale yellow. By day 2 xanthophores usually 

were present over the hindbrain, and by day 3 a third 

of the larvae had 1–2 over the forebrain. During the latter part of preflexion stage xanthophores proliferated 

dorsally and spread dorsolaterally on the head, and 

in flexion stage they were present over the forebrain, 

spread ventrolaterally to the level of the eyes and otic 

capsule, and spread posteriorly to myomeres 1–2. By 

early postflexion stage nearly the entire brain area and 

posteriorly to about myomere 3 was densely covered, a 

few xanthophores were scattered on the upper half of 

the opercular area, and a few were present on the snout 

and along the mesial ends of the maxillaries, premaxillaries, and dentaries. Pigmentation gradually increased 

in these areas. The opercular area and isthmus became 

silvery in pelagic juvenile stage.

Xanthophores formed on the trunk and tail during 

the latter part of preflexion stage. A row of irregularly 

spaced chromatophores formed along each side of 

the dorsal margin posteriorly to about the end of the 

melanophore series (myomere 23–25), with a few laterally, resulting in a bar in the vicinity of myomeres 
22

19–23. Some of the dorsal xanthophores flanked the 

melanophore row and others were interspersed among 

them; the majority on the trunk and flanking the melanophores were orange while most of those interspersed 

among the melanophores were yellow. Two rows of 

irregularly spaced xanthophores extended forward 

along the ventral margin from myomere 24–25 to anywhere between mid-tail and the first postanal myomere. 

Ventral xanthophores were predominantly yellow, and 

those in the tail bar were an ~ 50:50 mixture of orange 

and yellow. During notochord flexion pigmentation 

increased in all three areas, extending the full length of 

the tail ventrally, and spread laterally from the dorsum 

and ventrum, apparently beginning at the tail bar and 

progressing cephalad. A few xanthophores formed on 

the horizontal septum of the posterior half of the tail 

in flexion stage. In postflexion stage the tail bar broadened to myomeres 18–19 through 24–25. By pelagic 

juvenile stage xanthophore and melanophore patterns 

on the trunk and tail were essentially the same, except 

that xanthophores were scattered along the myosepta 

on the lower half of the tail where melanophores lacked. 

Xanthophores were dense in the saddles and bars.

Xanthophores formed dorsally on the gut at the end 

of preflexion stage and surrounded it except at the 

terminus of the hindgut in flexion stage. A few formed 

on the lower half of the cleithrum during postflexion 

stage. The abdominal area became silvery by pelagic 

juvenile stage.

The pectoral and caudal were the first fins to acquire 

xanthophores: during flexion stage a few (yellow) 

formed on the outer surface of the pectoral-fin base and 

proximally on the finfold, and 1–2 (orange) formed at 

the base(s) of the central caudal-fin rays. More formed 

on one or both surfaces of the pectoral-fin bases and 

proximally on some rays in postflexion stage. Xanthophores formed on the hypural margin in postflexion 

stage, and along all but the distal 10–25% of each principal caudal-fin ray by pelagic juvenile stage. Those on 

the dorsal and anal fins corresponded to melanophore 

patterns, but in the pelagic juvenile they extended a 

little farther distally on the soft-ray portion of the dorsal 

fin, and formed a stripe on the anal fin that extended 

from the middle of the second spine to the base of the 

next to last ray, with few accompanying melanophores. 

There were a few xanthophores proximally on the pelvic 

fins in pelagic juvenile stage.

Black-and-yellow rockfish 

(Sebastes chrysomelas)

Morphology Larval S. chrysomelas were 4.4–4.8 mm 

(mean 4.6 mm) at birth and completed yolk absorption 

3.5–5 days later at 4.6–5.4 mm (mean 5.1 mm). Neither 

notochord flexion nor development of the head spines 

and fin rays began before the last larvae died on day 9. 

The larvae were morphologically similar to S. atrovirens: 

moderately slender with rounded head and short snout, 

short preanal length, and slightly inflated, sac-like integument enclosing the trunk and anterior part of the 

tail (Table 9; Fig. 8). The larvae had 25–27 myomeres 

(97% with 26): 6–8 preanal + 18–20 postanal (68% with 

7 + 19, 26% with 6 + 20).

Pigmentation: Melanophores Larvae were moderately pigmented at birth (Fig. 8). Initially, the head was 

unpigmented except for 1–2 melanophores dorsally 

on the myelencephalon in 72% of the larvae. All larvae 

had a row of melanophores dorsally on the trunk and 

tail, usually 11–20 (mode 17) between myomeres 6–10 

and 22–24 (modes 7 and 9 through 23). The first 1–4 

melanophores in the series were more widely spaced 

than the others in 64% of the newly born larvae. In half 

of the newly born larvae the dorsal melanophores were 

in a single row; in the others the row was doubled at ≥ 1 

myomere(s), usually between myomeres 19–22. At birth 

some larvae (16%) had a melanophore on or near the 

horizontal septum on one side of the tail (usually the 

left) between myomeres 11–14; the others lacked lateral 

melanophores. About 45–61 (mean 53) melanophores 

were arrayed in a 1–3 melanophore-wide row (most with 

a 1–2 melanophore-wide row) ventrally on the tail, usually from myomere 7 through 23 or 24. The notochord 

tip was unpigmented. The pigment ventrally on the tail 

nearly always was continuous with that over the gut. The 

gut was heavily pigmented dorsally from near the level 

of the cleithra to the terminus. Ventrally, the gut was 

moderately pigmented, usually with 11–14 (mode 13) 

melanophores covering most of the mid-gut area and 

one on the hindgut adjacent to the anus. The gut was 

unpigmented laterally. 

Pigmentation changed only a little in the remaining 

days, except on the head. The proportion of larvae with 

melanophores over the myelencephalon increased to 

100% by day 4, but the number of melanophores remained 1–3 (modally 2). A melanophore or two formed 

over the cerebellum by day 4 in some larvae ≥ 5.3 mm 

(27%), and another 1–2 were added on each side of the 

cerebellum in a few larvae ≥ five days old (14%). One 

to three melanophores were present over the midbrain 

area in 24% of larvae > four days old. The number of 

melanophores ventrally on the tail increased slightly, 

to 47–69 (mean 58). One melanophore was present on 

the peritoneum at the anteroventral margin of the liver 

in 10% of the 3–4 day old larvae, and in 34% of older 

larvae. 

Pigmentation: Other chromatophores Xanthophore 

patterns at birth were not recorded. On day 4, xanthophores (yellow) were continuous on the dorsum from 
23

Table 9

Summary of measurements of black-and-yellow rockfish, Sebastes chrysomelas, given as percentages of body length in 1 mm 

size classes. For each measurement mean values are given above and ranges below; n = number of specimens.

Size range n PAL BD HL HW SnL ED P1L P2L

4.1–5.0 78 35 16 20 10 5 7 5 0

 33–38 14–18 19–22 9–11 4–6 6–8 4–5 

5.1–6.0 25 36 15 19 10 5 6 4 0

 34–37 14–16 19–20 9–10 4–6 6–7 4–5

the midbrain area to the end of the dorsal melanophore 

series (myomere 22–24) in 70% of the larvae, increasing 

to 80% by one week. In the remainder, the series was 

interrupted at myomeres 6–7 through 8–12. The dorsal 

xanthophores on the trunk and tail were irregularly 

spaced in two rows flanking the median melanophore 

row, with a few interspersed among the melanophores. 

No xanthophores were present on the gut at day 4, but 

by one week some had formed dorsally or dorsolaterally 

on the hindgut (present in 50%) and by day 9 ~ 90% 

of the larvae had 1–2 xanthophores on the peritoneum 

at or near the anterior margin of the liver. Most larvae 

(~ 90%) had a xanthophore near the center of the otic 

capsule, and at day 9 three-quarters had 1–2 xanthophores at or near the pharyngobranchial location. All 

larvae lacked ventral xanthophores. 

Discussion

Larval S. atrovirens, S. auriculatus, and S. chrysomelas are 

much alike, but exhibit some differences in xanthic and 

melanistic pigmentation, at least during the preflexion 

stage. Xanthophore patterns become increasingly alike 

between S. atrovirens and S. auriculatus after preflexion 

stage, and melanophore patterns become increasingly 

alike among most Pteropodus species after preflexion 

stage. Larval S. auriculatus lack dorsal xanthophores on 

the trunk and tail until late preflexion stage, while the 

other two have extensive dorsal xanthic pigmentation 

from birth. Larval S. chrysomelas lack lateral and ventral 

xanthophores on the tail for at least the first nine days, 

and S. auriculatus lack them until late preflexion stage, 

in contrast to S. atrovirens which have them ventrally, 

and often laterally, from birth. Larval S. atrovirens also 

have more extensive xanthic pigmentation on the jaws 

and gut than the other two. These differences suggest 

that xanthophore patterns could be useful for species 

identifications. However, xanthic pigmentation has not 

been documented for all the Pteropodus species and it 

is unknown how many have unique patterns. Preflexion-stage S. caurinus have a pattern much like that of 

S. atrovirens, but usually have a little xanthic pigment on 

the upper jaw (lacking in S. atrovirens before postflexion 

stage). During at least the first week larval S. rastrelliger

apparently lack xanthophores and thus resemble S. 

auriculatus, which have only a few xanthophores before 

late preflexion stage. Xanthophores rarely persist longer than a few days in the usual formalin or alcohol preservatives and their utility for identifying field-collected 

larvae currently is limited. (Attempts to stabilize the 

xanthic pigment in larval S. atrovirens using solutions of 

0.1–10.0% ionol in ethanol and in formalin were unsuccessful.) Nevertheless, documentation of xanthophore 

patterns may prove to be a valuable exercise if future 

research leads to a method for stabilizing this pigment 

in preserved specimens.

Melanophore patterns differ among larval S. atrovirens, S. auriculatus, and S. chrysomelas during the preflexion stage primarily in the location of the dorsal series, 

and in the number of melanophores and width of the 

ventral series on the tail. Dorsal melanophores on the 

trunk are rare in S. auriculatus but more common in 

the other two (~ 40–50% of specimens). Larval S. dalli

(Moser and Butler, 1981), S. caurinus (Stahl-Johnson, 

1985), S. carnatus (Moreno, 1993), S. rastrelliger (Moreno, 1993), and S. nebulosus (Kendall1) also usually have 

melanophores dorsally on some of the trunk myomeres. 

Larval S. auriculatus usually have fewer ventral melanophores on the tail than S. atrovirens and S. chrysomelas, 

averaging 30 in a single row; essentially the same pattern 

is displayed by larval S. dalli (Moser and Butler, 1981). 

Larval S. chrysomelas typically have the most ventral 

melanophores among the three, averaging 55 in a 1–3 

melanophore-wide series; essentially the same pattern is 

displayed by S. carnatus (Moser, 1967). Larval S. atrovirens are intermediate, with an average 45 melanophores 

in a 1–2 melanophore-wide series, and their range 

(28–62 melanophores) overlaps both S. auriculatus and 

S. chrysomelas. Larval S. rastrelliger initially have a ventral 

series like S. auriculatus, with 19–46 (mean 30) melanophores mostly in a single row, but the number increases 

and by 4–5 days after birth there are 48–61 (mean 52) in 

a 1–3 melanophore-wide series, much like S. chrysomelas. 
24

Figure 8

Black-and-yellow rockfish, Sebastes chrysomelas, preflexion stage. (A) 4.6 mm, day 1, lateral view; (B) 5.3 mm, day 6, lateral 

view; (C) 4.6 mm, day 1, dorsal view; (D) 5.3 mm, day 6, dorsal view; (E) 5.3 mm, day 6, ventral view.

A

B

C

D

E
25

Larval S. caurinus also initially have a predominantly 

single row of ~ 30–50 (mean 40) melanophores ventrally 

on the tail, but within a week the number increases to 

~ 35–60 (mean 45) in a 1–3 melanophore-wide series.

Intraspecies variation in pigmentation can be relatively large. For example, preflexion-stage larvae from 

one S. atrovirens had 28–49 (mean 42) ventral melanophores on the tail and 8–19 (mean 11) in the dorsal series, those from another had 37–52 (44) and 6–15 (11), 

respectively, and those from a third had 38–62 (48) 

and 4–15 (9), respectively (Table 10). In a single batch 

of S. auriculatus the number of dorsal melanophores 

on the trunk and tail in preflexion stage ranged from 

0 (in 2%) to 19. Preflexion-stage S. caurinus reared at 

SWFSC (March–April 1999) typically were more lightly 

pigmented, especially dorsally, than those described 

by Stahl-Johnson (1985), although the same batch 

also contained larvae as heavily pigmented. Marliave 

et al.3 described similar variation in a single batch of 

Table 10

Trunk and tail melanophores of preflexion-stage Sebastes larvae, subgenus Pteropodus. Under number of melanophores, the 

mean is given above and range below; for location of pigment, the myomere number at the modal location is given above 

and range below. SWFSC refers to larvae reared at Southwest Fisheries Science Center.

Dorsal Lateral Ventral

Species Source Number Location Number Location Number Location

S. atrovirens SWFSC 11 12–23 < 1 20 42 8–23

group 1 8–19 1–23 0–1 20 28–49 8–24

SWFSC 11 11–22 < 1 44 8–23

group 2 6–15 1–23 0–2 10–23 37–52 7–24

SWFSC 9 15–23 < 1 48 7–23

group 3 4–15 1–23 0–4 11–22 38–62 6–24

S. auriculatus SWFSC 13 11–22 1 20 30 9–23

 0–20 9–22 0–4 12–23 16–35 9–23

S. carnatus Moser, 1967 16 55

11–23 49–63

S. caurinus SWFSC 19 7–23 < 1 11 42 7–24

15–27 1–24 0–1 11–21 32–57 7–24

Stahl-Johnson, 1985 23

18–32

S. chrysomelas SWFSC 16 7, 9–23 < 1 10 55 7–24

 8–25 2–24 0–2 2–22 45–69 7–25

S. dalli Moser & Butler, 1981 14 2 38

 7–19 12–23 0–6 29–49 9–24

S. rastrelliger SWFSC 44 1, 7–23 1 12 57 8–23

26–55 1–24 0–3 2–24 43–82 7–24

S. nebulosus Kendall1 ca. 50 5–23 ca. 7–23

3 Marliave, J. B., D. I. Kent, and C. M. Brenton. 1997. Pigment polymorphism in sibling larvae of Sebastes caurinus. Abstr., 77th Ann. 

Meeting Amer. Soc. Ichthyol. Herpetol., Seattle, WA, 26 June–2 July 

1997. Available from University of Washington, College of Fisheries, Seattle, WA 98195.

S. caurinus, and we observed the same variability in 

S. chrysomelas.

Given the similar melanophore patterns among Pteropodus species and the variability within species, the 

instability of xanthic pigment in the usual preservatives, and the morphological similarity among species, 

it seems likely that unless a new method for preserving 

xanthic pigment is developed, reliable species identifications of field-collected larvae will not be possible using 

traditional morphological and pigment characters. 

However, because larval Pteropodus do have a common 

basic pattern of melanistic pigmentation during the 

preflexion stage (Fig. 9), identification of the subgenus 

may be possible. The principal elements of the pattern 

are the long dorsal melanophore row on the tail that 

often extends onto the trunk (Fig. 10), the long ventral 

melanophore row on the tail that originates at the last 

preanal or first postanal myomere (Fig. 11), and little 

or no pigment on the pectoral fins. Occasionally an 

individual may have a shorter dorsal row (occupying 

≤ 5–6 myomeres), or a ventral row that originates at 

the second postanal myomere. The basic pattern may 

be diagnostic for Pteropodus although larvae of some 

other species (e.g., squarespot rockfish S. hopkinsi: 
26

Figure 9

Preflexion-stage larvae of the Sebastes subgenus Pteropodus, lateral view. (A) Sebastes atrovirens, 4.6 mm; (B) S. auriculatus, 

5.8 mm; (C) S. carnatus, 4.3 mm (Moreno, 1993); (D) S. caurinus, 5.2 mm; (E) S. chrysomelas, 5.3 mm; (F) S. dalli, 6.2 mm 

(Moser and Butler, 1981); (G) S. maliger, ca. 5 mm (DeLacy et al., 1964); (H) S. nebulosus, 5.7 mm (Kendall1); (I) S. rastrelliger, 5.6 mm.

A

B

C

D

E

F

G

H

I
27

Figure 10

Preflexion-stage larvae of the Sebastes subgenus Pteropodus, dorsal view. (A) Sebastes atrovirens, 4.6 mm; (B) S. auriculatus, 

5.8 mm; (C) S. caurinus, 5.2 mm; (D) S. chrysomelas, 5.3 mm; (E) S. dalli, 6.2 mm; (F) S. rastrelliger, 5.6 mm.

A

B

C

D

E

F
28

A

B

C

D

E

F

G

Figure 11

Preflexion-stage larvae of the Sebastes subgenus Pteropodus, ventral view. (A) Sebastes atrovirens, 4.6 mm; (B) S. auriculatus, 

5.8 mm; (C) S. caurinus, 5.2 mm; (D) S. chrysomelas, 5.3 mm; (E) S. dalli, 6.2 mm; (F) S. nebulosus, 5.7 mm (Kendall1); 

(G) S. rastrelliger, 5.6 mm.
29

Taylor4) can display a similar pattern. During flexion 

and postflexion stages pigmentation of Pteropodus and 

several other species converge on a common pattern 

with melanophores on most or all of the dorsal margin, on the horizontal septum of the tail (especially 

posteriorly, often with a saddle or bar posteriorly), and 

posteriorly on the ventral margin of the tail. However, 

it may be possible to distinguish preflexion-stage larvae 

of the subgenus as a whole from other Sebastes larvae 

based on the Pteropodus pattern. Clearly, it is unlikely 

that every preflexion-stage Sebastes larva could be unequivocally assigned to Pteropodus or “other” categories: 

lightly pigmented Pteropodus would be excluded from 

that category and unusually pigmented non-Pteropodus

would be erroneously included. Probably ~ 75% of the 

preflexion-stage Pteropodus would be correctly identified 

and perhaps fewer than ~ 25% of non-Pteropodus larvae 

incorrectly classified based on larval pigmentation patterns. The likelihood of correctly classifying Pteropodus

larvae using pigmentation diminishes rapidly after the 

preflexion stage. Nevertheless, because ~ 90% of the larval Sebastes collected in standard oblique bongo net tows 

off California are preflexion stage (Moser, 1996b), if the 

pigmentation method for larval identification is validated and confidence bounds on the technique established, 

probably using molecular methods, it may be possible 

to generate an index of larval Pteropodus abundance as a 

fishery-independent measure of population trends for 

this group of nearshore rockfishes as a whole.

Acknowledgments

Many people were involved in collection of the rockfish 

broodstock that provided the larvae described here; 

we thank Cindy Taylor, Russ Vetter, Bill Cobb, Jason 

Stannard, Eric Lynn, Elaine Sandknop, Sherri Charter, 

Dave Griffith, Dave Ambrose, Amy Hays, Carol Kimbrell, 

Vince Buonaccori, Sean Narum, and the officers and 

crew of the NOAA research vessel McArthur. Broodstock 

fish were collected during the California Sea Grant 

Marine Ecological Reserves Research Program study 4-

M-N, conducted by Southwest Fisheries Science Center. 

Geoff Moser reviewed an early draft of the manuscript 

and provided many helpful comments.

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