NOAA Technical Report NMFS 116 November 1993

Description of Early Larvae of

Four Northern California Species of

Rockfishes (Scorpaenidae: Sebastes)

from Rearing Studies

Guillermo Moreno

u.s. Department of Commerce
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NOAA Technical Report NMFS 116

A Technical Report of the Fishery Bulletin

Description of Early Larvae of Four

Northern California Species of Rockfishes

(Scorpaenidae: Sebastes)

from Rearing Studies

Guillermo Moreno

November 1993

u.s. DEPARTMENT OF COMMERCE

Ronald H. Brown, Secretary

National Oceanic and Atmospheric Administration

D.James Baker, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

Rolland A. Schmitten, Assistant Administrator for Fisheries
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ii
Description of Early Larvae of Four Northern California Species

of Rockfishes (Scorpaenidae: Sebastes) from Rearing Studies

GUILLERMO MORENO

P. O. Box 450

Moss Landing Marine Laboratories

Moss Landing, CA 95039

ABSTRACT

About 72 species of Sebastes (Family Scorpaenidae) are found along the eastern Pacific

coast of North America, some of which are heavily exploited by both commercial and

sport fisheries. Because of the large number of species, the identification of early life

stages has progressed slowly. The objectives of this study were 1) to rear the larvae of

four species of rockfish (Sebastes mystinus, S. carnatus, S. atrovirens, and S. rastrelliger); and

2) to describe the larvae using morphometric measurements, pigmentation patterns,

and head spination.

Pigmentation was the most useful feature for identification purposes. Two general

patterns were found: 1) a short row of ventral midline melanophores on the tail, and

none or very little postero-dorsal pigmentation (S. mystinus); and 2) complete ventral

midline pigmentation on the tail, and anterior and postero-dorsal melanophores (S.

carnatus, S. atrovirens, and S. rastrelliger). With the exception of very early stages of S.

carnatus and S. atrovirens, these species can be readily identified. Morphometric proportions and head spination did not show major differences among species.

Because of the great similarities found among species in this genus, descriptions from

field studies are uncertain to some extent. Laboratory rearings, although difficult, can

at least provide early larvae from known species which allow precise identification as

well as an estimation ofvariability of characters (e.g., pigmentation) within and between

broods.

Introduction _

The genus Sebastes (Rockfishes; Family Scorpaenidae)

is an abundant and diverse group o£fishes along the

Pacific coast of North America (Love and Westphal,

1981) including about 72 species in 11 nominal subgenera (Kendall, 1991). The group is an important

part of the commercial and sport catches off California, Oregon, Washington, Canada, and Alaska (Moser

et aI., 1977; Gunderson and Lenarz, 1980;.- Lenarz,

1987). In California, rockfishes represent one third

of the recreational fish catch; S. mystinus, S. melanops,

and S. flavidus compose up to 30% of this proportion. Sebastes mystinus is also an important component ofthe commercial rockfish catch (Lenarz, 1987).

Species identification of early life history stages is

critical for systematics as well as for studies using

Present address: School of Biological Sciences, Zoology/ A08, The

University of Sydney, New South Wales 2006, Australia.

larval abundance for population estimates. In the

genus Sebastes, many larval characters are shared by

many closely related species, making determination

and evaluation of the importance of characters difficult (Moser et aI., 1977; Moser and Ahlstrom, 1978;

Barsukov, 1981; Kendall and Lenarz, 1987). Larval

rearing can provide essential identification information and permit an evaluation of variation in diagnostic characters oflarvae from different ages, broods,

and species. The relative importance of these diagnostic characters may then be evaluated.

Rockfishes are viviparous, extruding 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

yolk, but with well-developed eyes and mouths that

allow immediate feeding (Boehlert and Yoklavich,

1984) .

Partial descriptions are available for the larvae of

51 eastern Pacific rockfish species (Matarese et aI.,
2 NOAA Technical Report NMFS 116

1989); many important commercial and recreational

species remain undescribed. Early developmental

stages traditionally have been described from larvae

extruded in the laboratory and from preserved field

collected specimens (Morris, 1956; Moser, 1967, 1972;

Waldron, 1968; Westrheim, 1975; Moser et aI., 1977;

Moser and Ahlstrom, 1978; Richardson and Laroche,

1979; Laroche and Richardson, 1981; Moser et aI.,

1985). Also larvae have been reared from known

parents through the stage of caudal-fin formation

(Moser and Butler, 1981; Stahl-Johnson 1984, 1985;

Moser and Butler, 1987, Kendall 19891; Wold, 1990).

Efforts to rear larval Sebastes in captivity have met

with varying success. Japanese workers have raised

larvae of eight northwestern Pacific species, three of

these to the juvenile state (S. pachycephalus [Fujita,

1957; Siokawa and Tsukahara, 1961]; S. oblongus

[Fjuita, 1958]; and S. schlegeli [Hoshiai, 1977;

Kusakarai et aI, 1977]). However, only recently have

eastern Pacific Sebastes larvae been raised past yolk

absorption and, in some cases, to caudal-fin formation (Moser and Butler, 1981, 1987; Stahl-Johnson,

1985; Kendall, 19891; Wold, 1990). Factors responsible for difficulties in rearing northeastern Pacific

species may include their relatively small size at birth

«4.5 mm) and cool ambient water temperatures that

slow growth rates relative to western Pacific species

(Kendall and Lenarz, 1987).

The purposes of this study were 1) to rear larvae of

four rockfish species that are widely distributed along

the west coast of North America (Eschmeyer et aI.,

1983) and are associated with kelp beds or nearshore

rocky reef areas, or both, and 2) to describe their

early development. The species studied were blue (S.

mystinus) , gopher (S. carnatus) , kelp (S. atrovirens) ,

and grass (S. rastrelliger) rockfishes.

Methods

Collection

Gravid females of S. mystinus (n=4) and S. carnatus

(n=3) were collected at Stillwater Cove, Carmel Bay,

California, between December and May 1988 and

1989. Rockfishes were captured by SCUBA divers using hook-and-line or gill nets and transported to the

Monterey Bay Aquarium (MBA). Gravid females of S.

atrovirens (n=2) and S. rastrelliger (n=I) from the Kelp

Tank at the MBA were also used in this study. All

females were placed in rearing tanks until after parturition. Food (squid, krill, and anchovies) was oflKendall, A. W., Jr. Additions to knowledge of Sebastes larvae

through recent rearings. Dep. Commer., NOAA, Northwest and

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

fered to all females before parturition, but no feeding was observed.

Rearing

Rearing tanks with black walls and white conical bottoms held 400 L of continuous flowing (3-4 L/min)

of filtered (1 J.1m) seawater. The flowthrough system

was similar to that of Hughes et ai. (1974), differing

only in the direction of the flow, which in this study

was from top to bottom. At parturition, or soon after,

the females were removed from the rearing tanks,

and larval densities were reduced to <10 larvae/L (as

recommended by Stahl-Johnson, 1985) by siphoning

excess larvae until desired densities were reached.

Larvae were fed with a variety of food organisms.

Larvae up to one week old were fed 3-4 times per day

with the marine rotifer Brachionus plicatilis and wild

plankton. The plankton was collected daily in Elkhorn

Slough, Monterey County, CA, with a 30-J.1m net. All

items that passed through a 120-J.1m mesh were fed

to the larvae. Possible prey items observed included

rotifers, copepods, tintinnids, nauplii, trochophore larvae, and phytoplankton, mainly diatoms. Mter 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.1m 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., Isochrysis spp.,

and Duniolella spp.; Guillard, 1975) and were enriched daily with a mixture of fish oil (herring) and

egg yolk (1:1)2. The Artemia nauplii were enriched

with "Selco" (Artemia Systems N.V., Belgium), and

the wild plankton was treated with a fungicide-bactericide (1-10 ppm; "Prefuran," Argent Labs., Redmond,

WA) to reduce the possibility of bacterial contamination of the larval rearing tanks. During the first rearing season, water quality samples were taken weekly

and analyzed for nitrate, nitrite, phosphate, ammonia, and oxygen levels. Owing to consistently high

quality of the water system, these tests were later

discontinued. Continuous lighting was used to allow

24-hour feeding.

Descriptions

Larval fish were sampled (numbers depending on

availability) every four to five days for descriptive

work, photographs, and preservation. Larvae were

fixed initially in 5% buffered (sodium borate) forma2Refik Orhun, Hubbs Marine Research Center, San Diego, CA

92101. Pers. Commun.
Moreno: Description of Four Species of Rockfish Larvae 3

lin, and then transferred after 30 days to a dehydrating series of isopropanol (25-50%) and ethanol

(70%), allowing a week between each alcohol change.

Initially, measurements were taken before and after

fixation and dehydration to determine possible

shrinkage. No changes were noted so subsequently

larvae were measured only after preservation.

Morphometric and pigmentation analyses were conducted on dehydrated larvae for 0.5-mm SL size

classes within the available size range (e.g., 3.0-7.9

mm) for each species. 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) (Moser et aI., 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.

A modification of the pigment scheme of Kendall

and Lenarz (1987) of melanophore presence or absence at 26 loci was used. Pigmentation analysis of 33

loci (Fig. 1) 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 melanophores at these loci could

be counted with confidence. It was also possible to

count the number of melanophores in loci 22 and 23

for S. mystinus. The other loci had tightly packed

melanophores that made counts impossible in most

cases. Because of the large observed variability in

4 31 6 1117 18 19 33 20

I.... .

2' 3 ''''' ..... ''

29

28

10 12 13 14 16 15 30

....

@

: .....

7 9

8

21 27 22 23 24 25 26

Figure 1

Diagram of 33 melanophore positions found on Sebastes larvae. Positions are as follows (pectoral fin loci (7, 8, and 9) are

shown separately): Locus: 1 = symphysis of the lowerjaw; 2 = upperjaw; 3 = lateral aspect of the lowerjaw; 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 fin; 8 = blade of the pectoral

fin; 9 = margin of the pectoral fin; 10 = ventral aspect of the gut; 11 = dorsal aspect the gut; 12 = postero-dorsal surface of

the gut; 13 = postero-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 fin; 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 = internal locus above the dorsal surface the notochord from the 3rd to the 19th postanal myomere. (Modified

after Kendall and Lenarz, 1987.)
4 NOAA Technical Report NMFS 116

melanophore development, proportions of larvae by

size class with pigment at each locus were noted in

this study.

Common pigmentation sites in Sebastes larvae include the dorsal and lateral surfaces of the gut (loci

11, 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 lowerjaws

(loci 1-3, and 29), and caudal fin (loci 20, 25, and

26) (Kendall and Lenarz, 1987). Means, standard deviations, and ranges of melanophore numbers at these

loci were calculated for each 0.5-mm SL size class.

Descriptions were done with dissecting microscopes

(10-60x). An Olympus image analysis system (CUE

II) was employed for the morphometric measuremen ts. Illustrations conform to the guidelines in

Sumida et al. (1984); descriptive terminology and

methods follow Moser et al. (1977), Moser and

Ahlstrom (1978), Richardson and Laroche (1979),

and Kendall and Lenarz (1987). Statistical analyses were

not conducted because of the low number of independent samples (i.e., birthing females) that were used.

Results _

Descriptions of Larvae

The species described here had several common characters at birth that are shared with all Sebastes species

described to date: 1) pigmented eyes and functional

mouths; 2) an undifferentiated finfold surrounding

the tail; 3) pigmentation on the dorsolateral surface

of the gut (locus 11) and posterior surface of the

hindgut (locus 17); and 4) pigmen tation on the ventral surface of the tail (Morris, 1956; Moser, 1967,

1972; Waldron, 1968; Westrheim, 1975; Moser et aI.,

1977; Moser and Ahlstrom, 1978; Richardson and

Laroche, 1979; Laroche and Richardson, 1980,1981;

Moser and Butler, 1981, 1987; Moser et aI., 198.5;

Stahl-Johnson, 1985; Kendall and Lenarz, 1987; Wold,

1990). The extent of postanal midline pigmentation

helps to separate groups of species (DeLacy et aI.,

1964; Moser, 1967; Westrheim, 1975; Moser et aI.,

1977). In this study, the larvae of S. mystinus had

short ventral series of melanophores (loci 22 and

23), which are different from the longer, denser series (loci 21-23) characteristic of the other three

species (S. carnatus, S. atrovirens, and S. rastrelliger).

In addition, S. mystinus lacked, or had very little,

pigmentation on the dorsal portion of the tail (locus

19). This contrasted with the other three species

which had complete dorsal series (loci 18 and 19).

Sebastes mystinus-Larvae were obtained from three

females during January and February 1989. Two

broods were born; one with fully absorbed yolk and

the other with large (0.6-mm mean diameter) yolk

sacs. The latter brood had higher mortalities and the

larvae were often seen at the surface swimming on

their sides and occasionally upside down owing to

the buoyancy of the yolk sacs and oil globules. Larvae

from these two broods lived 10 and 11 days, respectively. A third female gave birth to larvae, some of

which survived 31 days. Some larvae from this brood

showed signs ofstarvation such as dorsoventrally compressed heads, "duck billed" jaws, and reduced body

depths. At 28 days the marine copepod Tigriopus

caliJornicus was introduced into the diet of this brood

to supplement the rotifer-brine shrimp nauplii-wild

plankwn diet. This copepod proved to be an unsuitable food item because of its large size and to its

aggregation along the sides of the tanks away from

the majority of the larvae.

The larvae ranged in SL from 3.2 mm at birth to

5.5 mm at day 31. The mean standard length at birth

was 3.8 mm (SD=0.2), with a mean of 4. 7 mm (SD=O.4)

at day 31 (Fig. 2). 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 (Fig. 3). This species had the largest

pectoral fins of any of the species described here.

Pigmen tation patterns of these larvae at birth were

different from the other three species (Table 1; Fig.

3). At birth, larvae had pigmentation on the dorsal

surface of the gut (locus 11) and anus (loci 16 and

17); the short ventral midline series (loci 22 and 23)

was composed of relatively few melanophores

(mean=15.4). By a mean length of 5 mm (day 21),

the majority of larvae had developed pigmentation

on the lower jaw (locus 1), forebrain (locus 4), nape

(locus 6; Fig. 4), otic capsule (locus 31), and cleithral

symphysis (locus 28). Melanophores were also beginning to appear posteriorly on the dorsal surface of

the tail (locus 19; Fig. 5) and on the margin and

bladt- of the pectoral fins (loci 8 and 9). The second

and third posterior preoperculars, and pterotic spines

were first observed on 4.7 mm mean standard length

larvae (31 days; e.g., Fig. 3C).

Sebastes carnatus-Three field-caught pregnant S.

carnatw gave birth in the rearing tanks during March,

ApriL and May 1989. One of these released larvae

with large yolk sacs (0.6-mm mean diameter) and

large quantities of oil. First feeding of these larvae

was observed after five days (4.0-mm 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
Moreno: Description of Four Species of Rockfish Larvae 5

8

+1 7

e

5

£ 6 btl

=qj

...:l

"C

I-< 5 <II

"C

=<II

... -t =<II

qj

3

0 10 20 30

Age (days)

40

S.mys/inus

-..-- S. carnQtus

---6- S. atrovirens

S. rasereWger

50 60

Figure 2

Mean standard length (± SE) vs. age of larvae of Sebasles myslinus (n=4), S. carnatus

(n=3), S. atrovirens (n=2), S. rastrelliger (n=1). n= number of females that gave birth.

Table 1

Proportions of Sebasles myslinus larvae with melanophores present at loci 1-33 (see Fig. 1 for loci description).

SL (mm) 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

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 (mm) 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n

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

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 parasite was probably introduced by

an infected female.

The range in standard length of these larvae was

3.1 mm at birth to 6.2 mm at 28 days. The mean

standard length of yolk-sac larvae was 3.4 mm

(SD=O.l) at birth and 5.8 mm (SD=0.2) at 28 days

(Fig. 2). The full-term larvae were 4.4-mm (SD=0.3)

mean standard length at birth. The pectoral fin

reached to just in front of the anus by the time a

mean standard length of 4.3 mm (day 13) was attained (Fig. 6).

The yolk-sac larvae showed little pigmentation

(Table 2; Fig. 6A). However, complete ventral (loci

21-23) and dorsal midline (loci 18 and 19) pigment

series were present (Fig. 5) along with a few melanophores scattered laterally on the yolk-sac and a prominent antero-lateral tail melanophore (locus 27). Melanophores on the dorsal surface of the gut (locus

11), an s (loci 16 and 17), and the antero- and

poster -dorsal aspect of the tail (loci 18 and 19) were

also quite frequent. All individuals in the 5.0-5.4 mm
6 NOAA Technical Report NMFS 116

Sebastes mystinus

c

@

.. " ..

'" ; .

..';..•. )\-. I.":."!

Figure 3

Sebastes mystinus larvae. (A) 4.2-mm SL preflexion larva; (B) 4.8-mm SL preflexion larva; (C) 5.6-mm SL

preflexion larva.

size class (day 13) had pigment on the jaw (locus 1),

forebrain (locus 4), nape (locus 6; Fig. 4), 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; Fig. 6C). Spination was first observed on larvae of 6.1-mm mean standard length

(25 days) with the appearance of the second and

third posterior preoperculars, pterotic, and parietal

spines (Fig. 6C).

Sebastes atrovirens-Larvae of S. atrovirens were obtained from two females removed from the MBA Kelp

Tank in May 1988 andJune 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.

Larvae from the full-term brood lived eight days;

some from the mixed brood lived for 30 days.

The range of larval standard length was between

4.0 mm at birth and 6.5 mm at day 30. The mean

standard length at birth was 4.3 mm (SD=0.2) with a

mean of 5.3 mm (SD=0.6) at 30 days (Fig. 2). The

pectoral fins extended to just in front of the anus by

the time a mean length of 4.5 mm (day 10) was

reached (Fig. 7).
Moreno: Description of Four Species of Rockfish Larvae 7

n

Mean -1- Range

-Standard Deviation

30 Locus 4

20 13 6 3

'"

CII

""III' -

00

~s=

ClIo. 5 .0 0

"II E c :> III 10 z- iIH CII

::E 21 6 5 59 24 3

1: ~III 32 19 :iIII :t:II

0

1 2 3 4 5 2 3 4 5 67 345 3 4 5 6 7 8 9 10

S. mys6nus S.carnatus S.atrovirens S. rastrelfiger

30 Locus 6

13 3

6

'" 15 I CIl 20 -

00

~IHf ~s=

CIl a. .0 0 7 5 E c

:> '" 10 z-CII 10 II 32 19 3 5924 ::E

i1~II I III 6 5

4 3821 I I

0 :l:I~

1 2 3 4 5 234 5 6 7 345 3 4 567 8 9 10

S. mys6nus S.carnatus S.atrovirens S. rastrelfiger

Species

Figure 4

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.

At birth, S. atrovirens larvae 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),

postero-dorsal midline of the tail (locus 19), and a

complete ventral midline series (loci 21-23) (Table

3; Fig. 7). With development, increasing proportions

of larvae showed pigmentation on the lower jaw (locus 1), forebrain (locus 4), nape (locus 6; Fig. 4),

anterior dorsal midline (locus 18; Fig. 5), and otic

capsule (locus 31) . With continued growth, pigmentation appeared on the posterior and postero-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 (see Pigmentation in Species Comparisons Section for explanation of differences). No head spine development was

apparent during the 30 days (Fig. 7D).
8 NOAA Technical Report NMFS IIG

n

I

- Range

Mean _ .. _Standard Deviation

30 Locus 18

13

VI

'" 20 00

.... .c "'a.

DOEc

:> til 10

z-;

o

21 4 36 6 5

... ... i" + +

7

15 6 3

5 272511.1 II

III 1

75

<II

'" 55 00

.... .c Cllc.

DOE c 35 :> til

z-'" 15

0

1 2 3 4 5

S. mystinus

4 36 21 6 5

++=m=.=-=

S. mystinus

2 3 4 5 6 7 3 4 5 3 4 5 6 7 8 9 10

S.camatus S.atrovirens S. rastrelliger

Locus 19

3

25 15 I

iIliU:

1 2 :3 4 5 6 7 3 4 5 3 4 5 6 -7 8 9 10

S.carnatus S.atrovirens S. raslrefliger

Species

Figure 5

Melanophore densities (mean number per lan'a) at locus 18 and locus 19 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.

Sebastes rastrelliger-One female S. mstrelligerremoved

from the MBA Kelp Tank in mid-February 1989 gave

birth on capture, but the larvae appeared to be fullterm because no yolk sacs or oil globules were present.

Some of these larvae lived up to 56 days.

Larval sizes ranged from 4.3 mm at birth to 7.7 mm

at day 52. The mean length at birth was 4.6 mm (SD=O.l)

and at day 56 was 6.6 mm (SD=0.3; Fig. 2). The pectoral

fins in this species were smaller than the other species,

reaching to -75% of the gut length (Fig. 8).

This was the most highly pigmented of the four

species described (Table 4; Fig. 8). 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

blfth to 20.0 and 18.7 at 7.5-7.9 mm (Fig. 4). 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
Moreno: Description of Four Species of Rockfish Larvae 9

Sebastes carnatus

A

:'-'.' :~·~"'7"';--W~.:-'"'I'7"7··;·"'·Y··::··

.--:~,:~.; ;~~." :~----

c ~'."" .'fil .. .

~. 7

Figure 6

Sebastes carnatus larvae. (A) 3.5-mm SL yolk-sac larva; (B) 4.3-mm SL preflexion larva; (C) 6.0-mm standard length

preflexion larva.

mm length was reached (Table 4). Melanophore density in the antero-dorsal midline series (locus 18)

increased from 3.6 to 15.7. The postero-dorsal midline series (locus 19) also increased, from a mean of

44 to >44 at 7.5-7.9 mm (Fig. 5). Because of the

dense arrangement of melanophores, it was not possible to quantify their exact number at this locus.

Distinctive postero-lateral tail pigment (loci 24 and

33) was present on most 5.0-5.4 mm larvae and

present in all 6.5-6.9 mm larvae. This pigmentation

was also seen on similar-sized S. carnatu5 larvae.

Spination was present on larvae of 6.3-mm mean

standard length at 29 days: the parietal, pterotic,

second and third anterior preoperculars, second,

third, and fourth posterior preoperculars, upper opercular, and postocular spines were present (Fig. 8C). By

7.4-mm mean standard length at 34 days the parietal,

pterotic, first, second, and third anterior preoperculars,

second, third, and fourth posterior preoperculars, upper opercular, and postocular spines were observed

(Fig. 8D). Hypural elements were developed on 7.4-

mm mean standard length larvae (34 days old).
10 NOAA Technical Report NMFS 116

Table 2

Proportions of Sebastes carnatus larvae with melanophores present at loci 1-33 (see Fig. 1 for loci description).

SL (mm) 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

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 10.

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 10.

4.5-4.9 0.1 0.0 0.0 0.1 0.0 0.8 0.0 0.0 0.0 0.8 10 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 (mm) 18 19 20 21 22 23 24 23 26 27 28 29 30 31 32 33 n

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 0.0 010 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 00 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

Table 3

Proportions of Sebastes atrovirens larvae with melanophores present at loci 1-33 (see Fig. 1 for loci description).

SL (mm) 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

4.0-4.4 0.1 0.0 0.0 0.1 0.0 0.1 0.0 O.V 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 00 0.0 1.0 1.0 0.1 0.1 0.1 0.1 1.0 1.0

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

SL (mm) 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n

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 0.1 0.0 0.0 32

4.5-4.9 0.4 1.0 0.0 1.0 1.0 1.0 0.0 o.n 0.0 0.0 0.3 0.0 0.0 0.3 0.0 0.0 19

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

Species Comparisons _

Morphometrics-Sebastes mystinus was characterized by

having a relatively longer snout (SNL/HL), and

deeper body (BD/SL) than the other species at similar sizes. In addition, this species had a larger head

(HL/SL) in relation to body length (Tables 5-8).

Although not measured, this species had the largest

pectoral fins. On larvae 4.8 mm or larger, the fins

extended behind the anus (Fig. 3).

Sebastes carnatus appeared to have a longer gut

(SA/SL) than S. rastrelliger for all size classes. It also

had a longer snout (SNL/HL) than S. atrovirens and

S. rastrelliger in the 4.0-6.4 mm ranges (Tables 6-8).

Sebastes atrovirens had a longer gut (SA/SL), than

the other species except S. carnatus at the 4.0-4.4

mm and 5.5-6.4 mm ranges. More notably, this species had large eyes (ED/HL) that may help to distinguish it from the very similar S. carnatus larvae (Tables

6 and 7).

Sebastes rastrelliger, along with S. mystinus, had a

relatively shorter gut (SA/SL), and deeper body (BD/

SL) compared with the other species. It also had

smaller eyes (ED/HL) than did S. mystinus and S.

rarnatus (Tables 5, 6 and 8).

Pigmentation-The pigment patterns which help to

differentiate S. mystinus from the other species described are I) 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 postero-dorsal midline pigment (10-
Moreno: Description of Four Species of Rockfish Larvae 11

Sebastes atrovirens

A

~~~:;==:::Z==~·~=::':~:;.'"

" -~~.: ""11': --.~-~-- __. .

\!)~.. .~"... --,"~~-~~ ..• ,;,.;~.~~,"::.>.> r - "".~"

_------ /

c

D ""~"":"""" 9""'·· "'~'-.

';;.' ...>...".

Figure 7

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.
12 NOAA Technical Report NMFS 116

Sebastes rastrelliger

c " •.. ~'-"~"""""''''"'/'''~'')~'''-:')'~''''7'~'V:''';:~~~~":~""~,,,,,~~,,,._~-----

'@~; .•_.....:,•.. '.'i":"""'.,~~ .• !)p")'" ... ,.. ...... """

. ..#---..:....;..;..~ .... ,.~~"'.,i._.,_ ..~.~~ ..:~- -:;,...~.:~:~:~~~:;:-~:.~~ '~.. ' .-. --.:::::

» ""'- .----_.

. ...... :"

'~,.,. -";~""""7"'~::~""., .;. -..-~:-,::-..::-,-'

:~' .....-:.....'.':::'. ··l··· . ..' ..•~:Y" .:: ;,;..;;;.,~ ...:\:';-;>-("':'-~\'

Figure 8

Sebastes rastrelliger larvae. (A) 4.6-mm SL yolk-sac 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.
Moreno: Description of Four Species of Rockfish Larvae 13

Table 4

Proportions of Sebastes rastrelliger larvae with melanophores present at loci 1-33 (see Fig. 1 for loci description).

SL (mm) 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17

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 0.4 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 (mm) 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 n

4.0-4.4 1.0 1.0 0.0 1.0 1.0 1.(; 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 09 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

Table 5

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; S L/HL=snout length/head

length; EDjHL = eye diameter/head length; BD/SL = body depth/standard length.

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

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) (3~.3-57.1) (25.0-50.0) (12.8-19.4)

4.0-4.4 34.9±1.7 20.6±l.i" 37.2±5.1 35.8±3.4 15.9±2.0 21

(31.7-39.5) (18.6-25.0) (25.0-454) (25.0-40.0) (12.1-19.5)

4.5-4.Q 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) (2 J.5-24.0) (41.6-.'';4.5 ) (33.3-36.3) (16.0-20.0)

cus 19; Fig. 5) 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 (Fig. 3).

Sebastes carnatus (Fig. 6) and S. atrol'irens (Fig. 7)

can be differentiated from S. mY5tinus (Fig. 3) by 1)

presence of a long ventral midline melanophore series (loci 21-23); and 2) a greater number of melanophores on the dorsal midline (loci 18 and 19; Fig.

5). They can be differentiated from S. rastrelliger (Fig.

8) by the overall more pronounced pigmentation on

the latter. Sebastes carnatus can be differentiated from

similarly pigmented S. atrovirens by the following characters: 1) presence (up to 0.9) of antero-Iateral tail

pigment (locus 27; Tables 2 and 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 mid-upper and mid-lower jaw

(loci 2 and 3; Table 2; Fig. 6).

Sebastes rastrelliger can be differentiated by the following patterns: 1) heavily pigmented head (loci 1-4

and 6; Figs. 4 and 8) with presence of melanophores
14 NOAA Technical Report NMFS 116

Table 6

Morphometric proportions of Sebastes carnatus 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.

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

3.0-3.4 40.5±2.6 15.9±2.2 24.6±7.9 41.2±5.0 9.S±4.5 16

(35.2-45.1) (11.7-21.2) (16.6-40.0) (33.3-50.0) 5.S-21.2)

3.5-3.9 39.7±1.9 17.S±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.S) (2S.5-60.0) (5.5-17.9)

4.0-4.4 3S.S±2.0 lS.7±1.6 34.2±6.0 37.0±5.S ] 3.3±5.0 59

(34.S-42.S) (15.9-23.2) (25.0-50.0) (25.0-50.0) (2.0-19.0)

4.5-4.9 36.S±1.4 IS.5±l.l 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.S±1.5 20.5±1.0 33.6±4.1 34.6±4.3 17.5±6.3 10

(35.S-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.S 24.2±1.7 32.2±3.9 33.3±2.4 20.0±1.4 7

(39.6-44.S) (22.0-27.1) (26.6-37.5) (30.7-35.7) (17.5-22.0)

6.0-6.4 42.S±1.S 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)

Table 7

Morphometric proportions of Sebastes atrovirens 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.

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

4.0-4.4 3S.l±2.1 lS.S±2.1 31.0±6.5 37.S±6.1 14.l±1.9

(34.0-43.9) (13.9-23.S) (12.5-44.4) (22.2-57.1 ) ( 11.3-1S.6)

4.5-4.9 3S.0±2.7 19.4±2.2 2S.3±5.4 3S.l±5.7 13.4±2.1

(33.3-44.S) (15.2-2S.4) (20.0-40.0) (25.0-50.0) (10.6-1S.3)

5.0-5.4 39.9±2.S 21.S±3.1 27.6±7.3 39.1±2.2 14.3±1.6

(35.S-45.0) (1S.S-29.4) (16.6-40.0) (36.3-41.6) (11.3-16.6)

5.5-5.9 41.S 21.S 25.0 50.0 14.5

6.0-6.4 42.2±1.S 24.7±3.1 24.l±4.6 42.7±7.4 17.5±2.1

(40.9-44.2) (22.9-2S.3) (21.4-29.4) (35.2-50.0) (16.3-19.9)

n

50

3S

9

3

on the nares (locus 32); 2) complete, heavily pigmented dorsal midline series (loci 18 and 19; Fig. 5)

from birth; and 3) external and internal pigmentation on the postero-Iateral portion of the tail (loci 24

and 33) in larvae >5.0 mm (Table 4; Fig. 8). The

extent and concentration of ventral midline pigment

(loci 21-23) is similar to that of S. carnatus and S.

atrovirens.

Discussion

The traditional size-series method may not be practical for all species of Sebastes, because many fieldcaught Sebastes larvae smaller than 8-10 mm cannot

be identified with confidence (Kendall and Lenarz,

1987). The very early stages of S. carnatus, S. atrovirens

(this study), S. chrysomelas (Wold, 1990), S. auriculatus
Moreno: Description of Four Species of Rockfish Larvae 15

Table 8

Morphometric proportions of Sebastes rastrelliger 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.

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

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) (11.1-16.7)

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)

and S. caunnus (Stahl:Johnson, 1985) show great similarities and are difficult to differentiate from each other.

Of the 51 species of eastern Pacific Sebastes larvae

previously described, only nine descriptions have been

based on reared specimens (Moser and Butler, 1981,

1987; Stahl-Johnson, 1985; Kendall, 1989; Wold, 1990)

mainly because of the difficulty of rearing larvae past

yolk-sac to juvenile stage. Such difficulties arise from

the relatively small size, early developmental level at

birth, feeding problems and importantly, slow larval

growth rates shown for eastern Pacific species (Moser

and Butler, 1987).

A major concern in rearing Sebastes is the need for

healthy full-term larvae (Westrheim, 1975). Most information on the early larval stages of rockfishes

comes from descriptions of pre-extrusion larvae. However, pigment patterns are not well established in

yolk-sac Sebastes larvae, making these descriptions

unreliable for the identification of more advanced

stages (Westrheim, 1975). Because of the use of

SCUBA-aided capture techniques, premature birth

was alleviated for the nearshore Sebastes species described here.

The months of parturition observed for two (S.

mystinus, S. carnatus)' of the four species were similar

to those reported by Wyllie-Echeverria (1987). There

is no published information on parturition periods

of S. atrovirens or S. rastrelliger. Sebastes species seem to

have two major seasons of larval extrusion, winter

and spring-summer (Phillips, 1964). Sebastes mystinus

and S. rastrelliger gave birth in the winter season (N0-

vember-March), and S. atrovirens in the spring-summer season (April-July). Sebastes carnatus overlapped

both seasons with March-May parturitions.

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. The data presented here for the other

three species are still 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 these 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 1) grouping by age is a slow and expensive

process in field studies and 2) 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
16 NOAA Technical Report NMFS 116

most important determinant in the timing of metamorphosis of the starry flounder, Platichthys stellatus.

It is relatively easy to identifY two (S. mystinus, S.

rastrelliger) of the four species described here by pigmentation patterns. However, morphometric measurements did not yield differences useful for positive identification of species. Sebastes mystinus had

larger pectoral fins than the other species, but because ofthe difficulty in measuring the unpigmented

pectoral fins of the other species it was not possible

to compare them. Sebastes camatus and S. atrovirens are

very similar and pigmentation alone will not be enough

to identifY all the stages of either species. These larvae show the following morphometric differences: S.

camatus has a longer snout (SNL/HL) and deeper

body (BD/SL), whereas S. atrovirens has a larger eye

diameter (ED/HL). Parturition times and range information are also important to decide which species may

be present in the plankton at a certain time and place.

Two distinct pigmentation patterns can be discerned among the larvae described here: 1) a short

row of ventral melanophores and no, or very little,

postero-dorsal pigmentation (S. mystinus); and 2) complete ventral pigmentation (e.g., all three loci pigmented) and anterior and postero-dorsal melanophores (S. camatus, S. atrovirens, and S. rastrelliger).

Sebastes mystinus had pigmentation on the head and

on the margin and blade of the pectoral fins. Sebastes

camatus and S. atrovirens had similar melanophore

patterns in the very early stages; S. camatus differed

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 more melanophores per locus than any ofthe other species. Among

these four species, S. rastrelliger was unique in having

melanophores on the nares. The internal posterolateral pigmentation on the tail of S. rastrelligerand S.

camatus is also presentin S. caurinus and S. auriculatus

(Stahl-Johnson, 1985) and may commonly be found

in older larvae.

Other approaches to the identification of Sebastes

larvae include capture and rearing of planktonic larvae (Kendall, 1991) and electrophoretic techniques

(Seeb and Kendall, 1991). However, these methods

have limitations as well. 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), whereas autolytic

tissue decomposition may start within two to three

minutes after sampling (O'Connell, 1980; SetzlerHamilton et aI., 1987) possibly affecting electrophoretic results.

Some investigators (Morris, 1956; DeLacy et aI.,

1964; Moser, 1967; Efremenko and Lisovenko, 1970;

Westrheim, 1975) have expressed doubts about the

possibility of identifYing field-caught Sebastes larvae

because of small interspecific differences and large

intraspecific variability. However, some species whose

larvae have already been described are identifiable

in spite of these problems. Researchers need to describe in more detail the degree ofvariability of characters commonly used for species identification (e.g.,

pigmentation).

Although difficult, rearing of Sebastes species can

provide not only reliable descriptions but also useful

information on various life history aspects. In the

case of closely related species, this technique could

partially eliminate the uncertainty associated with

descriptions from field samples. In addition, culturing can provide much needed data on larval growth,

age, and behavior.

Acknowledgments _

This work is the result of research sponsored in part

by NOAA, 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. This

grant was awarded to G. M. Cailliet and V. Loeb.

Additional support was given to the author by the

David and Lucile Packard Foundation.

lowe many thanks to my committee members, G.

M. Cailliet, V. Loeb, and P. Roe, for their support

and interest in my work, and for critically reviewing

this manuscript; to L. Wold, who worked with me

setting up this project and solving numerous problems; to M. M. Yoklavich for reading the manuscript

and helping with head spine identification; and to L.

McMasters, who drew the larvae and helped many

times with drafting and photographic work. Food

rearing techniques were vastly 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 MBA. This manuscript was greatly improved by the comments from

two anonymous reviewers. Finally, I would like to

thank the faculty, staff, and students of Moss Landing

Marine Laboratories for their extraordinary help and

encouragement.

Literature Cited

Ahlstrom, E. H.

1959. Vertical distribution of pelagic fish eggs and larvae

offCalifornia, and Baja California. U.S. Fish. Wildl. Serv.,

Fish. Bull. 161:107-146.
Moreno: Description of Four Species of Rockfish Larvae 17

Barsukov, V. V.

1981. A brief review of the subfamily Sebastinae. J.

Ichthyol. 21:1-26.

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.

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. 2:51-67.

Efremenko, V. M., and L. A. Lisovenko.

1970. Morphological features of intraovarian and pelagic

larvae of some Sebastodes species inhabiting the gulf of

Alaska. In P. A. Moiseev, (ed.), Soviet fisheries investigations in the northeastern Pacific, Part. V, p. 267-

286. Izv. Tikhookean. Nauchno-Issled. Inst. Morsk. Rybn.

Khoz. Okeanogr. (VNIRO) 70. [Transl. from Russ.: avail.

U. S. Dep. Com mer., Natl. Tech. Inf. Serv., Springfield,

VA 22161, as TT71-50127.]

Eschmeyer, W. N., E. S. Herald, and H. Hammann.

1983. A field guide to Pacific Coast Fishes ofNorth America.

Peterson Field Guide Series 28. Houghton Mifflin Co.,

Boston, 336 p.

Fujita, S.

1957. On the larval stages of a scorpaenid fish, Sebastes

pachycephalus nigricans (Schmidt). Jpn. J. Ichthyology

6:91-93. (In Japanese with English summary).

1958. On the egg development and larval stages of a viviparous scorpaenid fish, Sebastes oblongus Gunther. Bull.

Jpn. Soc. Sci. Fish. 24:475-479. (In Japanese with English

summary).

Guillard, R. R. L.

1975. Culture of phytoplankton for feeding manr,e

invertebrates. In W. L. Smith and M. H. Chanley (eds.),

Culture of marine invertebrate animals, p. 29-60. Plenum Press, New York and London.

Gunderson, D. R.. and W. H. LenarL.

1980. Cooperative survey of rockfish and whiting resources

offCalifornia, Washington, and Oregon, 1977. Mar. Fish.

Rev. 42(2):1.

Hoshiai, G.

1977. Larvae and juveniles of the scorpaenid fish, Sebastes

schlegeli .lap. J. Ichthyology 24:35-42. (In Japanese with

English summary.)

Hughes, J. T., R. A. Shleser, and G. Tchobanoglous.

1974. A rearing tank for lobster larvae and other aquatic

species. The Progressive Fish-Culturist 36:129-132.

Kendall, A. W.Jr.

1991. Systematics and identification of larvae and juveniles of the genus Sebastes. Env. BioI. Fish. 30:173-170.

Kendall, A. W. Jr., and W. H. Lenarz.

1987. Status of early life history studies of northeast Pacific

rockfishes. In Proceedings, international rockfish symposium, October 1986, Anchorage, Alaska, p. 99-

128. Alaska Sea Grant Rep. 87-2, Univ Alaska, Fairbanks.

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 Fish.

Exp. Station Monthly Rep. Vol. 34(10) Supplement, 11 p.

Laroche, W. A., and S. L. Richardson.

1980. Development and occurrence oflarvae and juveniles

of the rockfishes Sebastes Jlavidus and Sebastes melanops

(Scorpaenidae) off Oregon. Fish. Bull. 79:901-924.

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. flavidus, and S.

melanops. Fish. Bull. 79:231-257.

Lenarz, W. H.

1987. A history of California rockfish fisheries. In Proceedings, international rockfish symposium, October 1986,

Anchorage, Alaska, p. 35-41. Alaska Sea Grant Rep. 87-

2, Univ Alaska, Fairbanks.

Love, M. S., and W. V. Westphal.

1981. Growth, reproduction, and food habits of olive rockfish, Sebastes serranoides, off central California. Fish. Bull.

75:412-416.

Matarese, A. C., A. W. KendallJr., D. M. Blood, and B. M. Vinter.

1989. Laboratory guide to the early life history stages of

Northeast Pacific fishes. OAA Technical Report MFS

80,650 p.

Morris, R. W.

1956. Early larvae of four species of rockfish,

Sebastodes. Calif. Fish and Game 42:149-153.

Moser, H. G.

1967. Reproduction and development of Sebastodes

paurispinis and comparison with other rockfishes offsou thern California. Copeia: 773-797.

1972. Development and geographic distribution of the rockfish, Sebastes macdonaldi (Eigenmann and Beeson, 1893),

family Scorpaenidae, off southern California and Baja

California. Fish. Bull. 70:941-958.

Moser, H. G., and E. H. Ahlstrom.

1978. Larvae and pelagic juveniles of blackgill rockfish,

Sebastes rnelanostomus, taken in midwater trawls off southern California and Baja California. J. Fish. Res. Board.

Can. 35:981-996.

Moser, H. G., andJ. L. Butler.

1981. Description of reared larvae and early juveniles of

the calico rockfish, Sebastes dallii. Calif. Coop. Ocean.

Fish. Invest. Rep. 22:88-95.1987. Descriptions of reared

larvae of six species of Sebastes. In W. H. Lenarz and D.

R. Gunderson (eds.), Widow rockfish. proceedings of a

workshop, p. 19-29. NOAA Tech. Report NMFS 48.

Moser, H. G., E. M. Sandknop, and D. A. Ambrose.

1985. Larvae and juveniles of Aurora rockfish, Sebastes auro-rrl, from off California and Baja California. In A. W.

KendallJr. and.J. B. Marliave (eds.), Descriptions of early

life history stages ofselected fishes, p. 65-76. Can. Tech.

Rep. Fish. Aquat. Sci. 1359.

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 Helicolenus from

other oceans. NOAA Tech. Rep. NMFS Cire. 402, 71 P

O'Connell, C. P.

1980. Percentage of starving northern anchovy, Engraulis

rnordax, 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. 126,70 p.

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.

Richardson, S, L., and W. A. Laroche.

1979. Development and occurrence of larvae and juveniles

of the rockfishes Sebastes cramen, Sebastes pinniger, and
18 NOAA Technical Report NMFS 116

Sebastes helvomaculatus (Family Scorpaenidae) off

Oregon. Fish. Bull. 77:1-41.

Seeb, L. W., and A. W. KendallJr.

1991. Allozyme and DNA polymorphisms: new approaches

to the identification of larvae of genus Sebastes. Env.

BioI. Fish. 30:191-201.

Setzler-Hamilton, E. M., 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. Am. Fish.

Soc. Symp. 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. WorksJpn., 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 175, Rome, 100 p.

Stahl-:Johnson, K. L.

1984. Rearing and development of larval Sebastes caurinus

(copper rockfish) and S. auriculatus (brown rockfish) from

the northeastern Pacific. M. S. thesis, Univ. Washington, Seattle, 217 p.

1985. Descriptive characteristics of reared Sebastes caurinus

and S. auriculatus larvae. In A. W. Kendall Jr. and J. B.

Marliave (eds.), Descriptions of early life history stages of

selected fishes, p. 65-76. Can. Tech. Rep. Fish. Aquat.

Sci. 1359.

Sumida, B. Y., B. B. Washington, and W. A. Laroche.

1984. Illustrating fish eggs and larvae. In H. G. Moser et

al. (eds.), Ontogeny and systematics of fishes, p. 33-

35. Spec. Publ. No.1, Am. Soc. Ichthyol. Herpetol.

Waldron, K. D.

1968. Early larvae of the canary rockfish, Sebastodes

pinnigl'1". 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. Board. Can. 32:2399-

2411.

Wold, L.

1990. A practical approach to the description and identification of Sebastes larvae. M. S. thesis, Moss Landing Marine Laboratories, California State Univ., Hayward, 88 p.

Wyllie-Echeverria, T.

1987. Thirty-four species of California rockfishes: maturity

and seasonality of reproduction. Fish. Bull. 85:229-250.