NOAA Technical Report NMFS Circular 402

mm

Guide to the Identification
of Scorpionfish Larvae
(Family Scorpaenidae) in tlie
Eastern Pacific With
Comparative Notes on Species
of Sebastes and Helicolenus
From Other Oceans

H. Geoffrey Moser, Elbert H. Ahlstrom, and
Elaine M. Sandknop

April 1977

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

NOAA TECHNICAL REPORTS
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NOAA Technical Report NMFS Circular 402

_^0^M0S^

Guide to the Identification
of Scorpionflsh Larvae
(Family Scorpaenidae) In the
"^^^^^ Eastern Pacific With

Comparative Notes on Species
of Sebastes and Helicolenus
From Other Oceans

H. Geoffrey Moser, Elbert H. Ahlstrom, and
Elaine M. Sandknop

April 1977

U.S. DEPARTMENT OF COMMERCE

Juanita M. Kreps, Secretary

National Oceanic and Atnnospheric Administration

Robert M. White, Administrator

National Marine Fisheries Service

Robert W. Schoning, Director

For Sale by ihe Superintendent of Documents. U.S. Government Printing Office
Washington, DC. ^0402 Stock No. 003-020-00128-9

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publication.

CONTENTS

Page

Introduction 1

Materials and methods 2

Summary of larval characters of Scorpaenidae with emphasis on those important in identifying genera ... 3

Keys to larvae of eastern Pacific scorpaenid genera 7

Description of life history series 7

Sebastes Cuvier 8

Sebastes paucispinis (Ayres), Figure 5 9

Sebastes macdonaldi (Eigenmann and Beeson), Figure 7 16

Sebastes jordani Gilbert, Figure 9 18

Sebastes leuis (Eigenmann and Eigenmann), Figure 11 22

Sebastes cortezi (Beebe and Tee-Van), Figure 13 22

Sebastes sp. — Gulf of California Type A, Figure 15 26

Sebastes marinus (Linnaeus), Figure 17 26

Sebastes viuiparus (Kr«fyer), Figure 20 31

Sebastes fasciatus (Storer), Figure 21 33

Sebastes capensis (?) (Gmelin), Figure 21E 35

Sebastes — northwestern Pacific species. Figure 22 36

Heliocolenus Goode and Bean 38

Helicolenus dactylopterus (Delaroche), Figure 23 38

Sebastolobus Gill 40

Sebastolobus altivelis Gilbert, Figures 24 and 25 43

Sebastolobus alascanus Bean, Figure 26 46

Trachyscorpia Ginsburg 46

Scorpaenodes Bleeker 48

Scorpaenodes xyris (Jordan and Gilbert), Figures 27-29 50

Scorpaena Linnaeus 54

Scorpaena guttata Girard, Figure 31 55

Scorpaena Type A, Figure 33 57

Pontinus Poey 60

Pontinus Type A, Figure 35 61

Pontinus Type B, Figure 37 64

Ectreposebastes Garman 67

Ectreposebastes imus Garman, Figure 39 67

Acknowledgments 69

Literature cited 69

Figures

1. Larvae of Sebastes (subgenus Sebastomus) removed from pregnant females and cultured to the point

of yolk exhaustion 10

2. Larvae of Sebastes at stage of yolk exhaustion 11

3. Larvae of Sebastes at stage of yolk exhaustion 12

4. Larvae of Sebastes at stage of yolk exhaustion 13

5. Developmental series of Sebastes paucispinus 14

6. Stations at which larvae of Sebastes paucispinis were collected by CalCOFI plankton surveys during

1953 and 1956 15

7. Developmental series of Sebastes macdonaldi 17

8. Stations at which larvae oi Sebastes macdonaldi were collected during 4 yr (1953, 1960, 1965, and 1966)

of CalCOFI plankton surveys 18

9. Developmental series of Sebastes jordani 19

10. Stations at which larvae of Sebastes jordani were collected by CalCOFI plankton surveys during 1966 21

11. Developmental series of Sebastes levis 23

12. Stations at which larvae of Sebastes levis were taken during 4 yr (1951, 1955, 1968, and 1969) of Cal-
COFI plankton surveys 24

13. Developmental series of Sebastes cortezi 25

iii

14. Stations in the Gulf of California at which larvae of Sebastes cortezi were taken on CalCOFI cruises
during 1956 and 1957 26

15. Larvae of Sebastes Gulf of California Type A 27

16. Stations in the Gulf of California at which larvae of Sebastes Gulf of California Type A were taken on
CalCOFI cruises during 1956 and 1957 28

17. Developmental series of Sebastes marinus 29

18. Relation of snout-anus length to body length in developmental stages of Sebastes spp 30

19. Relation of pectoral fin length to body length in developmental stages of Sebastes spp 30

20. Larvae of Sebastes viviparus 32

21. Larvae of Sebastes fasciatus and a larva of Sebastes capensis 34

22. Larvae of Japanese species of Sebastes 37

23. Larvae of Helicolenus dactylopterus 39

24. Larvae of Sebastolobus spp 41

25. Developmental series of Sebastolobus altivelis 42

26. Developmental series of Sebastolobus alascanus 47

27. Larvae of Scorpaenodes xyris 50

28. Larvae of Scorpaenodes xyris 51

29. Developmental series of Scorpaenodes xyris 52

30. Stations at which larvae of Scorpaenodes xyris were taken on CalCOFI cruises during 1956 and 1957

and EASTROPAC expeditions during 1967 and 1968 54

31. Larvae of Scorpaena guttata 56

32. Stations at which larvae of Scorpaena guttata were taken on CalCOFI cruises during 1966 58

33. Larvae of Scorpaena Type A 59

34. Stations at which larvae oi Scorpaena Type A were taken on CalCOFI cruises in the Gulf of California
during 1956 and 1957 60

35. Developmental series of Pontinus Type A 62

36. Stations at which larvae and pelagic juveniles of Pontinus Type A were collected on CalCOFI cruises
during 1956 and 1957, EASTROPAC expeditions during 1967 and 1968, and Scripps Tuna Ocean-
ography cruises during 1958 and 1959 64

37. Developmental series of Pontinus Type B 65

38. Collections of larvae and pelagic juveniles of Pontinus Type B from various sources 67

39. Developmental stages of Ectreposebastes imus 68

40. Collections of larvae of Ectreposebastes imus from EASTROPAC expeditions, Scripps Tuna Ocean-
ography cruises, Scripps Institution of Oceanography fish collection, and the University of Hawaii . . 69

Tables

1. Charactersof the four subfamilies of scorpaenid fishes in the eastern Pacific 3

2. Meristics for scorpaenids included in this guide 4

3. Depth of pectoral fin base and length of pectoral fin relative to body length for larvae of seven scor-
paenid genera, expressed as mean percentage and range for three larval stages 5

4. Length (mm) at hatching, notochord flexion, transformation into pelagic juveniles, and known lengths

of pelagic juveniles of scorpaenid fishes included in this guide 6

5. Mean number and range of midline melanophores on tail portion of body of 33 species of Sebastes . . 9

6. Measurements (mm) of Sebastes paucispinis larvae 15

7. Meristics from cleared and stained larvae of Sebastes paucispinis 15

8. Mean lengths (mm) of Sebastes paucispinis larvae collected on cruises of the California Cooperative
Oceanic Fisheries Investigations during 1953 and 1956 16

9. Measurements (tnm) of larvae and pelagic juveniles of Sebastes macdonaldi 16

10. Meristics from cleared and stained larvae of Sebastes macdonaldi 18

11. Mean lengths (mm) of Sebastes macdonaldi larvae collected on cruises of the California Cooperative
Oceanic Fisheries Investigations during 1953 and 1960 18

12. Measurements (mm) of larvae of Sebastes jordani 20

13. Meristics from cleared and stained larvae of Sebastes jordani 20

14. Relative abundance and frequency of occurrence of Sebastes jordani in CalCOFI collections for 1966 21

15. Standardized numbers (A), mean length in millimeters with standard deviation (B), and size range in
millimeters of Sebastes jordani larvae taken on CalCOFI cruises of 1966 22

16. Measurements (mm) of Sebastes levis larvae 22

17. Measurements (mm) of larvae of Sebastes cortezi 24

iv

18. Mean length (mm) with total standardized number of larvae of Sebastes cortezi and S. Type A in
plankton collections from the Gulf of California 24

19. Measurements (mm) of larvae of Sebastes sp. (Gulf of California Type A) 26

20. Measurements (mm) of larvae of Sebastes marinus 28

21. Measurements (mm) of larvae of Sebastes viviparus 31

22. Morphometric comparison of larvae of three species of Sebastes from the North Atlantic 32

23. Measurements (mm) of larvae of Sebastes fasciatus 33

24. Measurements (mm) of larvae of Sebastes capensis 35

25. Measurements (mm) of larvae of Helicolenus dactylopterus 40

26. Measurements (mm) of Sebastolobus spp. larvae 43

27. Measurements (mm) of Sebastolobus altivelis 44

28. Meristics from cleared and stained larvae and juveniles of Sebastolobus spp. and Sebastolobus altivelis 45

29. Measurements (mm) of larvae and juveniles of Sebastolobus alascanus 48

30. Meristics from cleared and stained larvae and juveniles of Sebastolobus alascanus , 49

31. Measurements (mm) of larvae of Scorpaenodes xyris 53

32. Meristics from cleared and stained larvae of Scorpaenodes xyris 53

33. Measurements (mm) of larvae of Scorpaena guttata 55

34. Numbers of occurrences and standardized numbers of larvae of Scorpaena guttata taken by CalCOFI

in 1966 57

35. Measurements (mm) of larvae of Scorpaena Type A 58

36. Measurements (mm) of larvae of Pontinus Type A 61

37. Measurements (mm) of larvae of Pontinus Type B 64

38. Measurements (mm) of larvae of Ectreposebastes imus 67

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

H. GEOFFREY MOSER, ELBERT H. AHLSTROM, and ELAINE M. SANDKNOP'

ABSTRACT

Developmental stages of 51 species or forms of scorpionfishes are described and illustrated in
this identification guide. Thirty -eight are from the eastern Pacific and represent six of the eight scor-
paenid genera known from that region — Sebaatea, Sebastolobus, Scorpaenodea, Scorpaena, Pontimia,
and EctrepoBebaatee. Sebaatea is the most thoroughly treated; developmental series of six species from
the eastern Pacific are described and illustrated; pigment patterns of early larvae of 33 species are
given and 23 of these are illustrated. Larval series of three North Atlantic species of Sebaatea are de-
scribed and illustrated as is a series of Sebaatea from off Chile; in addition, the published information
on eight northwestern Pacific species is summarized and discussed in relation to the eastern Pacific
and Atlantic species. The other genera are represented by one or two species. Since larvae of the
eastern Pacific species of Helicolenua were not available, a larval series of B. dactylopterua from the
Atlantic are described for comparative purposes. Larvae of the eighth eastern Pacific scorpaenid
genus, Trachyacorpia, are unknown.

Two dichotomous keys to the eastern Pacific genera are included, one for the early larval stages
up to the initiation of notochord flexion and one for postflexion larvae. In the text, a summary of the
literature and definitive characters is followed by the descriptive accounts of the species. Each species
account contains a literature summary, description, set of illustrations, and information on dis-
tribution and abundance.

INTRODUCTION

The scorpionfishes are among the most important
groups of shore fishes in the eastern Pacific. The ap-
proximately 100 species of eastern Pacific scorpionfishes
are distributed among eight genera — Sebastes,
Helicolenus, Sebastolobus, Scorpaenodes, Scorpaena,
Pontinus, Ectreposebastes, and Trachyscorpia. About
two-thirds of the species belong to the single genus,
Sebastes, which has undergone an extensive radiation in
the northeast Pacific. These rockfishes or rockcods
provide important commercial and recreational fisheries;
one species alone, Sebastes alutus, yielded 283,000 met-
ric tons to trawlers in a single year (Gulland 1970) and, in
California, rockfishes comprise one-half of the total num-
ber of sport fish taken annually by recreational anglers
(Young 1969).

The abundance of Sebastes is reflected in their
prominence in the plankton collections of the California
Cooperative Oceanic Fisheries Investigations (CalCOFI).
They are live-bearers and, at birth, the 4- to 5-mm larvae
rise to the surface waters where they may be sampled by
plankton nets (Moser 1967a, b). As a group, they usually
rank third or fourth in total abundance among all larvae
collected annually by the CalCOFI plankton survey
(Ahlstrom 1961). The task of identifying Sebastes larvae

'Southwest Fisheries Center, National Marine Fisheries Service,
NOAA, La Jolla, CA 92038.

to species is a difficult one because of the large species
complement of the genus; however, complete larval
series of some important commercial species have been
described (Moser 1967a, 1972, and in this guide). The
most productive technique has been to remove late-stage
embryos or prenatal larvae from identified pregnant
females and to raise these in the laboratory to a point
where they develop their specific pattern of
melanophores. These larvae can then be used to identify
developmental series of larvae from plankton collec-
tions.

During the investigation of Sebastes larvae, con-
siderable knowledge of the larvae of other eastern Pacific
scorpaenid genera has accumulated. One of these,
Sebastolobus, inhabits northern waters and its larvae are
found commonly in CalCOFI plankton samples from the
California Current. The larvae and highly distinctive
juvenile stages of S. altivelis and S. alascanus have
recently been described (Moser 1974). The other eastern
Pacific genera are primarily tropical-subtropical in dis-
tribution, with the exception of Helicolenus which occurs
off southern Chile. The larvae of these warm water
genera {Pontinus, Scorpaena, and Scorpaenodes) are en-
countered in the CalCOFI collections off southern Baja
California and in the Gulf of California as well as in
plankton collections from expeditions such as EAS-
TROPAC, and each has a character or group of characters
which distinguishes it from all other eastern Pacific
genera. The genus Ectreposebastes is primarily

equatorial in distribution and the larvae of £. imus were
taken on a few EASTROPAC stations.

It is the purpose of this paper to present information
and illustrations (including keys) for the identification of
the larvae of eastern Pacific scorpaenids to genus and, for
each genus, to describe and illustrate the larval stages of
the species for which larvae are known.

Sebastes is the most extensively treated. In addition to
summaries of the earlier descriptions of S. paucispinis
and S. macdonaldi, we include new descriptions of larvae
of S. jordani, S. levis, S. cortezi, and S. sp. (Gulf of
California Type A). Also, we describe larval series of
three North Atlantic species — S. marinus, S. uiviparus,
and S. fasciatus. Our purpose in doing this is to sup-
plement the already substantial literature on larvae of
these species with heretofore unreported characters and
to provide a basis for comparison of larvae of Atlantic
and Pacific Sebastes. Larval series of a Sebastes from off
Chile and eight species from Japan are described and il-
lustrated to allow comparison of northeastern Pacific lar-
vae with those from the Southern Hemisphere and
western Pacific. Essentially we are treating Sebastes on a
worldwide basis.

Since larvae of the eastern Pacific representative of
Helicolenus, H. lengerichi, were not available, we in-
clude a description of the Atlantic species, H. dactylop-
terus, so that larvae of Helicolenus may be compared
with those of other eastern Pacific genera. Many of the
characters presented in this description have not been
previously reported.

Larvae and juveniles of Sebastolobus have been
thoroughly described (Moser 1974) and are briefly sum-
marized here. The larvae of Trachyscorpia are unknown
but if their relationships are with Sebastolobus as Esch-
meyer (1969) suggests, then they probably will resemble
the larvae of Sebastolobus but with lower counts of most
meristics.

Life history stages of Scorpaenodes have not been de-
scribed and we give a detailed account of larvae and early
juveniles of the single eastern Pacific species, S. xyris.
The literature contains life history accounts of three
Mediterranean and North Atlantic species of Scorpaena
and descriptions of the eggs and newly hatched larvae of
a single eastern Pacific species, S. guttata (see literature
review for the genus). We include descriptions of a com-
plete larval series of S. guttata and of another Scorpaena
not identifiable to species. Larvae of Pontinus have not
been previously described and we describe larvae of two
forms of this genus. Two larval specimens of Ec-
treposebastes imus have been briefly described (Esch-
meyer and Collette 1966) and we describe a complete
developmental series of this species for the first time.

Seven of the eight genera of scorpaenids that occur in
the eastern Pacific also occur in the Atlantic. According
to Eschmeyer (1969, 1971) who reviewed the scorpion-
fishes of the Atlantic Ocean, 55 of the 60 species (i.e.,
92't) of Atlantic scorpaenid fishes belong to these seven
genera. Since there is a strong similarity between the
scorpaenid fauna of the two oceans at the generic level,
the guide provides a starting point for identification of

scorpaenid larvae in the tropical and subtropical Atlan-
tic.

In addition to the genera treated in this guide, the
literature contains relatively few other life history de-
scriptions of scorpaenid fishes. We are aware of pub-
lications on only three other genera: Pterois (P. lunulata.
Mito and Uchida 1958; Mito 1963), Inimicus {I.
japonicus, Fujita and Nakahara 1955), and Sebastiscus
{S. marmoratus, Tsukahara 1962). We are hopeful that
our guide will stimulate further investigations on the
developmental stages of this large and diverse family.

Relationships among the eastern Pacific genera can be
shown by their groupings into subfamilies. We are follow-
ing the classification presented by Matsubara (1943) and
amended for the subfamily Setarchinae by Eschmeyer
and Collette (1966) and Sebastolobinae by Eschmeyer
(1969). The eight genera of eastern Pacific scorpaenids
belong to four subfamilies, as follows:

Subfamily Sebastinae

Sebastes Cuvier

Helicolenus Goode and Bean
Subfamily Sebastolobinae

Sebastolobus Gill

Trachyscorpia Ginsburg
Subfamily Scorpaeninae

Scorpaenodes Bleeker

Scorpaena Linnaeus

Pontinus Poey
Subfamily Setarchinae

Ectreposebastes Garman

Among the primary characters for separation of adult
scorpaenid fishes into subfamilies are the shape of the
2nd suborbital (3rd infraorbital) bone, whether this bone
is fastened to the preopercle or remains free, and the
presence or absence of 3rd and 4th suborbital bones. The
suborbital sensory canal, which runs backward from the
preorbital to the 2nd suborbital in all scorpaenids has a
branch extending through the 3rd and 4th suborbital
bones to the 5th suborbital bone in some scorpaenids.
The latter branch is developed in eastern Pacific scor-
paenids that retain the 3rd and 4th suborbital bones.
Characters associated with the suborbital bones are sum-
marized for the eastern Pacific genera and subfamilies in
Table 1 together with meristic characters, type of
reproduction, and presence or absence of a gas bladder in
juveniles and adults.

MATERIALS AND METHODS

Specimens used in this study were obtained from a
variety of sources. The CalCOFI plankton collections
from the California Current region provided most of the
larval specimens of Sebastes and Sebastolobus. CalCOFI
collections from the southern end of the station pattern
off Baja California and from the Gulf of California
provided larvae of Sebastes, Scorpaena, Pontinus, and
Scorpaenodes. Larvae of the latter three genera were also

Table 1. Characters of the four subfamilies of scorpaenid fishes in the Eastern Pacific. Counts
listed are typical ones and occasionally encountered atypical counts are given in parentheses.

Subfamily

Genera

Suborbital bones

Suborbital
sensory
canal

Gas
bladder

Type

of

reproduction

Vertebrae

Dorsal
spines

Dorsal
rays*

Anal
rays*

Caudal

principal

rays

Caudal

branched

rays

2nd

3rd

4th

Sebastinae

Sebastes

Tapering to
a point, not
attached to
preopercle

Present

Present

Has exten-
sion from
2nd sub-
orbital to
5th sub-
orbital

Present

OvoviviDarous,
internal fer-
tilization,
larvae spawn-
ed

26-28
(E.Pac.)

29-32
(N.Atl.)

XIII-XIV
(E.Pac.)

XIV-XVI
(N.Atl.)

11-17

III
5-11

8+7

6+6

Helicolenus

Narrowed
posteriorly,
firmly at-
tached
preopercle

Present

Present

Same as
above

Absent

Probably
ovoviviparous
but eggs in
gelatinous
mass

25

XII
(XI-XIII)

11-13

111,5

8t7

6+5

Sebastolobinae

Sebastolobus

Rounded
posteriorly,
firmly at-
tached to
preopercle

Present

Present

Same as
above

Absent

Oviparous
eggs in
gelatinous
masses

28-31

XV-XVII

8-10

III
(4)5

8+7

6+5

Trachyscorpia

Not known

Present

Present

Same as
above

Absent

Same as above

25-26

XII

8-9

111,5

8+7

6+6

Scorpacninae

Scorpaena

Pontinus

Scorpaenodes

Uniformly
broad, or
becoming
wider pos-
teriorly,
firmly at-
tached to
preopercle

Absent
Absent

Absent

Present
Present
Absent

Lacks
exten-
sion to
5th sub-
orbital

Absent
Absent
Present

Oviparous
eggs in
gelatinous
masses
(Scorpaena)

24

24

24-(25)

XII

XII
XIII

8-10

8-9

9-11

III
5(6)

III
5(6)

III
5(6)

8+7
8+7
8+7

6+5
6+5
6+5

Setarchinae

Ectreposebastes

Same as
above

Absent

Absent

Same as
above

Absent
or

rudi-
mentary

Oviparous

24

XII

9-10

III

5(6)7

8+7

6+6

•Last soft ray of doral and anal fins is a double ray and is counted as a single ray.

found in pleinkton collections of the wide-ranging EAS-
TROPAC expeditions (Ahlstrom 1971, 1972) and in col-
lections from cruises of the Scripps Tuna Oceanography
Research (STOR) group (Blackburn et al. 1962). Pelagic
and benthic juveniles were obtained largely from ich-
thyological collections of the Scripps Institution of
Oceanography (SIO) and the Los Angeles County
Museum (LACM). Larvae and juveniles of Sebastes
marinus, S. viviparus, and Helicolenus dactylopterus
were obtained from the collections of the Dana Ex-
peditions at the University Zoological Museum, Copen-
hagen. A series of Sebastes fasciatus larvae from the
American coast between lat. 38°52'N and lat. 44°22'N
were supplied by the National Marine Fisheries Service
(NMFS) Laboratory at Narragansett, R.L, and a collec-
tion of intraovarian larvae from this species was ob-
tained from the NMFS Laboratory, Woods Hole, Mass.
Methods of analyzing meristic and morphometric
development follow Moser (1967a, 1972, 1974), however
some of the terms used in this study require further ex-
planation. The larval stage of teleosts can be divided
naturally into three substages based on the state of
development of the caudal fin. The substage beginning
with the termination of the yolk-sac stage and ending
with the initiation of notochord flexion we call preflex-
ion. The substage from the initiation of notochord flexion
to its completion, when the posterior edge of the hypural
plate arrives at a vertical position, we term flexion. From
this substage to the initiation of transformation is termed
postflexion. These substages provide a practical means
of comparing larval development in different taxa.

The change from larva to juvenile is termed transfor-
mation or transition. The juvenile stage may also be sub-
divided. Many scorpaenid species have a pelagic juvenile
substage that is marked by a distinctive pigment pat-
tern of bands and/or saddles and often by structural
features such as enlarged pectoral fins. When pelagic
juveniles become benthic juveniles there is usually an
abrupt change in pigment pattern and morphometry
(e.g., Sebastolobus altivelis). In species which remain
pelagic for much of their juvenile stage (e.g., Sebastes
jordani) there is no abrupt change in pigment or mor-
phometry.

In discussing distribution and abundance we use the
term standardized number of larvae. This means the ab-
solute number of specimens per tow adjusted to the num-
ber under 10 m- of sea surface (Kramer et al 1972).

SUMMARY OF LARVAL CHARACTERS OF

SCORPAENIDAE WITH EMPHASIS

ON THOSE IMPORTANT IN

IDENTIFYING GENERA

The most salient characteristic of scorpionfish larvae is
their elaborate head spination. Li addition to marked
development of spines on the preopercle, a feature shared
with many families, the larvae develop a pair of crestlike
parietal ridges that terminate in a single or double spine.
This parietal spination differentiates scorpaenid larvae
from those of most other teleost families. Larvae of the
Triglidae, a related family, also have parietal crests, but

these have scalloped margins and eire supported by rod-
like structures. The parietal crests in scorpaenids
typically have a serrate margin and have a single
strengthening rod at the posterior terminus.

In Sebastes the paired parietal spines on top of the
head develop in larvae 6.0 to 7.0 mm in length, and by
the midlarval period each projects backward from a ser-
rated parietal ridge. Later in the larval period, a smaller
nuchal spine usually develops adjacent and posterior to
each parietal spine. In Sebastolobus and Scorpaenodes,
the nuchal spines form soon after the parietals and rapid-
ly become larger and more prominent than the parietals.
This difference in size of the nuchals relative to the
parietals is especially useful in separating larvae of
Sebastes from those of Sebastolobus.

Spines also form on a number of head bones in addi-
tion to the parietals, e.g., the pterotics, frontals, nasals,
and circumorbitals (Moser 1972). Head spination is best
developed on postflexion larvae and is a primary charac-
ter in the key to postflexion larvae. Although they begin
to form on late preflexion larvae, head spines are seldom
differentiated enough to be used for identifying leirvae to
genus before flexion is completed.

As in other families, meristic characters are essential
in identifying larvae of scorpionfish to genus and species.
The most fundamental meristic character is the number
of myomeres, since it is the earliest to appear and is
equivalent to the number of vertebrae that will develop.
The temperate or boreal genera {Sebastes, Sebas-
tolobus, Helicolenus) have higher counts than the
tropical-subtropical genera {Scorpaena, Scorpaenodes,
Pontinus, Trachyscorpia, Ectreposebastes) as sum-
marized in Table 2.

Characters of the fins, including numbers of spines and
rays, relative size, shape, and pigmentation are in-
dispensable in identifying scorpaenid larvae. The pec-
torals are the first to appear and are usually the first in
which ossified rays are formed, followed by ossification of
principal caudal rays, rays and spines in the pelvics, dor-

sal and anal fins, and finally procurrent caudal rays. Os-
sified rays begin to form in the pectoral fins prior to
notochord flexion; they form sequentially from the dor-
sal to the ventral margin. The 15 principal caudal rays
and pelvic rays (I, 5) form during flexion, whereas the
dorsal, anal, and procurrent caudal rays begin to ossify
during late flexion and early postflexion stages. The com-
plete complement of pectoral rays forms within a 3-mm
size range whereas the pelvic, dorsal, and anal fins form
within a 2-mm size range; only the formation of procur-
rent caudal rays is extended over a greater range of
length.

The number of pectoral rays is not particularly useful
in distinguishing among most genera of scorpaenids.
Most genera dealt with in this report have 16 to 20 rays.
The only striking exceptions are one species of Sebas-
tolobus with 21 to 24 rays and Trachyscorpia with about
22 to 23 rays.

The size, shape, and pigment pattern of the pectoral
fins are useful in distinguishing among genera and
species of scorpaenids (see key to genera for postflexion
larvae). Especially useful is the depth of the fin base,
which is narrowest in Sebastes, widest in Ec-
treposebastes, and of intermediate depths in the other
five genera. Values for this previously unutilized charac-
ter are summarized by species and stage of development
in Table 3.

There is a marked range in the proportional size of the
pectoral fin among the genera included in this study.
Larvae of Ectreposebastes have huge fan-shaped pec-
torals which extend posteriad to the base of the caudal
fin; they increase in relative length during development
to a maximum of one-half the body length (Table 3). The
fan-shaped pectorals in Scorpaenodes are only slightly
smaller than in Ectreposebastes, attaining a maximum
of 45"^ of the body length. The pectorals in Sebastolobus
and Pontinus are moderately large but differ strikingly in
shape, being fan-shaped in the former and aliform in the
latter. In Scorpaena they are fem-shaped and moderate in

Table 2. Meristics for scorpaenids included in this guide.

Vertebrae

Dorsal

Spinous

Rays

Dorsal
Soft
Rays

Anal
Soft
Rays

Pectoral
Rays

Caudal Rays |

Principal

Branched

i^rocurrent I

Sup.

Inf.

Sebastes jordani
" levis

macdonaldi
" paucispinis

27
26
26
26

XIII
XIII
XIII
XIII

13-16
12-13
13-14
13-15

8-11
6-7
7-8
8-10

19-22
17-18
18-20
15-16

8+7
8+7
8+7
8+7

6+6
6+6
6+6
6+6

13-14

11
13-14
12-13

13-14

11
13-14
12-14

Sebastes cortezi
capensis

26
26

XIII

xin

11-12
12-14

5-6
5-6

17-19
18-20

8+7
8+7

6+5
6+6

10-11

11-12

Sebastes marinus
" viviparus
" fasciatus

30-32
29-31
30-31

XIV-XVI
XIV-XVI
XIV-XVI

12-15
13-14
12-15

7-9
6-8
6-8

18-20
17-19
17-20

8+7
S+7
8+7

6+6
6+6
6+6

11-12
?

7

11-12
?

7

Helicolenus dactylopterus

25

XII

11-13

5

17-20

8+7

6+5

11

10-12

SebastolQbus alascanus
altiwelis

30-31
28-29

XV-XVII
XV-XVI

9-10
8-10

(4)5
5

21-22
22-24

8+7
8+7

6+5
6+5

8-10
6-8

8-9
7-8

Trachyscorpia sp.

25

XII

9

6

22-23

8+7

6+6

5

6

Pontinus Type A
" Type B

24
24

XII

XII

8-9
9

5(6)
5

18-19
17-19

8+7
8+7

6+5
6+5

6
5-7

6-7
5-7

Scorpaena guttata
■' Type A

24
24

XII
XII

8-10
8

5-6
5

17-19
19-20

8+7
8+7

6+5
6+5

6-8
5

7-9
5

Scorpaenodes xyris

24-25

XIII

9-11

5-6

16-19

8+7

6+5

4-5

4-5

Ectreposebastes imus

24

XII

9-10

5-7

18-20

8+7

6+6

6-7

6-7

Table 3. Depth of pectoral fin base and length of pectoral fin relative to body length for larvae of
seven scorpaenid genera, expressed as mean percentage and range for three larval stages.

Species

Number
specimens

Depth

of pec

toral fin base

Length of pectoral

fin

1

Preflexion

Flexion

Postflexion

Preflexion

Flexion

Postflexion |

X

range

X

range

X

range

x

range

x

range

x

range

Sebastes paucispinis

21

9.0

8-10

9.0

8-10

7.2

5- 8

16.4

11-21

27.2

25-31

35.5

34-37

macdonaldi

26

-

-

-

-

-

-

8.4

6-10

13.0

12-15

19.3

15-26

jordani
levis

23

6.7

6- 7

7.8

7- 8

7.8

7- 9

6.6

6- 7

8.4

7-10

14.7

11-20

12

8.0

-

12.8

12-14

11.7

11-12

9.0

-

34.9

24-46

43.3

41-45

cortezi

17

9.0

7-10

11.8

11-12

10.8

10-12

8.2

7- 9

10.6

9-12

17.5

11-21

Gulf Type A

13

7.8

5- 9

12.3

11-13

12.8

11-14

8.7

6-10

13.3

11-16

16.8

14-20

capensis

8

14.7

8-13

14.0

12-16

12.7

10-17

10.0

9-11

15.5

12-19

23.7

18-27

marinus

22

6.7

4- 8

8.2

8- 9

8.3

7- 9

6.0

3- 7

9.9

8-13

16.8

14-19

viviparus

14

7.5

6- 9

9.6

8-11

-

-

7.3

6- 9

9.9

8-12

-

-

fasciatus

14

6.8

6- 9

8.7

8- 9

8.7

8- 9

6.2

6- 7

8.0

6-10

15.3

13-18

Sebastolobus alascanus

40

11.0

6-15

15.8

13-19

14.4

13-16

12.7

7-17

22.2

19-25

29.0

26-31

altivelis

48

11.0

6-15

15.8

13-19

18.3

15-20

12.7

7-17

22.2

19-25

35.0

28-39

Helicolenus dactylopterus

16

11.0

9-12

12.5

12-13

12.4

10-14

11.4

11-12

14.8

13-16

18.0

16-20

Scorpaenodes xyris

36

15.0

8-18

18.7

17-20

18.0

16-21

22.2

10-28

34.2

29-41

39.0

36-45

Scorpaena guttata

16

13.8

8-16

16.0

15-17

15.9

14-17

15.4

7-22

20.8

19-22

21.7

18-23

Type A

18

13.8

12-15

14.6

13-16

17.3

16-18

12.2

10-15

16.2

15-18

20.7

19-23

Pontinus Type A

22

15.3

15-16

15.5

15-16

14.3

12-16

20.4

18-19

26.5

26-27

30.7

27-36

Type B

4

-

-

17.5

17-18

16.0

14-18

-

-

33.5

33-34

27.0

24-30

Ectreposebastes imus

12

20.4

19-22

22.0

-

21.0

19-22

32.6

24-38

38.0

-

44.4

37-49

relative length and in Helicolenus they are small and
rounded. In Sebastes there is a great variety in relative
size and shape among the numerous species. Also, dif-
ferent species attain maximum relative pectoral length
at different developmental stages (Table 3).

Patterns of melanistic pigmentation on the pectorals
are useful in identifying larvae to genus and species as
demonstrated in the key to preflexion larvae. The pat-
terns are too diverse to be summarized here and are best
treated in the succeeding sections.

Anal fin ray counts are not of particular value in dis-
tinguishing among most genera of scorpaenids. The usual
anal fin count is III, 5 (occasionally III, 4 or in, 6) in
Sebastolobus, Trachyscorpia, Helicolenus, Scorpaenodes,
Scorpaena, and Pontinus. In Ectreposebastes the count
is typically III, 6 with a range of 5 to 7 rays. The number
of anal rays has a greater range among species of
Sebastes. The typical counts for eastern North Pacific
species of Sebastes are HI, 6 and 7, however some species
have slightly higher ray counts and S. jordani has III, 8 to
11 anal rays. The second of the three anal spines is usual-
ly longer than the third and heavier in juvenile and adult
specimens. It should be noted that the third anal spine is
late in differentiating. When first formed in larvae, it has
the appearance of a soft ray. It is not ossified as a spine
until the early juvenile stage; however, we have desig-
nated the third element in the anal fin as a spine in all of
our counts of larvae. The change in the third anal ele-
ment to a spine is gradual. Even in a cleared and stained
series it is sometimes difficult to determine just when it
becomes a spine; however, the total count of spines plus
rays in the anal fin remains constant. For example,
whether the anal count for larvae of Sebastolobus is
given as H, 6 or III, 5, the total count remains 8. An iden-

tical problem with regard to the change-over of the third
element of the anal fin from a ray to a spine is encountered
in larvae of Mugil (Anderson 1957, 1958), and striped
bass, Morone saxatilis (Mansueti 1958). The anterior two
spines of the anal fin are supported in secondfiry associa-
tion by a single massive, elongated pterygiophore, pos-
sibly the product of the fusion of the first two; the re-
maining anal spine and the anal rays ase each supported
by a pterygiophore and are in secondary association with
the following pterygiophore (except the last ray). The
pterygiophore count is two less than the total count of
anal spines and rays.

The number of dorsal fin spines is a particularly useful
character (Table 2). The commonest count is XU, found
in Scorpaena, Pontinus, Helicolenus, Trachyscorpia, and
Ectreposebastes. Species of Sebastes in the eastern
North Pacific normally have XHI dorsal spines, as does
Scorpaenodes. North Atlantic species of Sebastes have
XIV to XVI dorsal spines, whereas Sebastolobus has XV
to XVII. The number of dorsal soft rays is usually 9 (8 to
10) in Sebastolobus, Trachyscorpia, Scorpaena, Pon-
tinus, Ectreposebastes, and Scorpaenodes, 11 to 13 in
Helicolenus, and 11 to 16 in Sebastes. The total number
of dorsal fin rays and spines is 20 to 22 in Pontinus,
Trachyscorpia, and Scorpaena, 21 or 22 in Ec-
treposebastes, 22 to 24 in Scorpaenodes, 23 to 25 in
Sebastolobus, and 24 to 31 in Sebastes.

The last spine in the dorsal fin of all the above scor-
paenids appears first as a soft-ray and later ossifies as a
spine, just as in the third spine of the anal fin. The an-
terior two spines of the dorsal fin are supported in second-
ary association by a single wide (fused) pterygiophore.
In eastern Pacific scorpaenids this pterygiophore lies
between neural spines two and three and is always ac-

companied in this space by a second pterygiophore,
which carries a single spine (the third). The succeeding
several pterygiophores alternate one on one with neural
spines. The first and second neural spines lie close
together and often point forward, in contrast to the diver-
gent angle between neural spines two and three neces-
sary to accommodate the anterior two pterygiophores. In
several genera a single, short predorsal bone was ob-
served immediately before the first neural spine; this
bone is more readily seen on cleared and stained
specimens than on radiographs.

The number of principle caudal rays (those supported
by hypurals and the parhypural according to our defini-
tion) in most scorpaenid fishes is 8 -I- 7 = 15. The number
of branched rays is either 6 -H 6 (Sebastes, Ectre-
posebastes) or 6 -I- 5 (Sebastolobus, Helicolenus, Scor-
paenodes, Scorpaena, Pontinus). Hence in these scor-
paenid genera there are two unbranched principal rays
associated with the upper lobe of the caudal fin, and
either one or two associated with the lower lobe — one un-
branched if the branched ray count of the lower lobe is
six, two unbranched if the branched ray count is five.
The principal rays of both lobes of the caudal fin are sup-
ported exclusively by hypural bones (we include the
parhypural as a hypural). The primitive condition,
which we have observed in Ectreposebastes imus, is the
retention of three superior and three inferior hypurals
(including the parhypural). Among the genera included
in this guide, Scorpaenodes and Pontinus retain the
primitive complement of three superior hypurals, but
have the inferior complement reduced to two. In the
other four genera there are two superior and two inferior
hypurals. The primitive complement of three epurals is
retained by all scorpaenids studied. Only one pair of
uroneurals was observed on specimens examined. The
neural spine on the vertebral centra immediately an-
terior to the ural is markedly reduced or lacking. The
haemal spine on this vertebra is autogenous, as is the
haemal spine on the vertebra immediately anteriad.

Since changes in body shape during larval develop-
ment are illustrated and are emphasized in the descrip-
tions, we will discuss them only briefly here. Newly
hatched larvae of all scorpaenids studied are more slen-
der, often markedly more slender, than are later larval
stages. Larvae of Sebastes and Helicolenus are usually
more slender in later larval stages than are those of the
other genera. The eastern Pacific Sebastes with the slen-
derest body is S. jordani; in postflexion larvae the
average relative body depth is less than 25% standard
length (range of 22 to 26''p). Relative body depth for post-
flexion larvae of other species ranges from 30 to 37%.
Average relative body depth for postflexion specimens in
the other genera is as follows: Sebastolobus, 35 and 41%
for its two species; Scorpaenodes, 39%; Scorpaena, 38 to
40% for two species; Pontinus, 39 and 42% for two species;
and Ectreposebastes up to 55%.

A knowledge of the sizes at initiation of major develop-
mental events such as hatching, notochord flexion, and
transformation aids in identifying scorpaenid larvae.
There is a major dichotomy between Sebastes and the

other genera in the size at hatching — Sebastes hatches at
3.8 to 7.5 mm body length and the other genera hatch at
1.8 to 2.8 mm (Table 4). Sebastes larvae are also longer
at notochord flexion (6 to 12 mm) than the tropical
genera Scorpaenodes, Scorpaena, Pontinus, and Ec-
treposebastes (4 to 6 mm) and the other temperate-
boreal genera, Sebastolobus and Helicolenus, are inter-
mediate (6 to 8 mm). Size at transformation from larva
to pelagic juvenile is highly variable among genera and
species of scorpaenids (Table 4).

Also essential for identification of scorpaenid larvae is
the locality of collection. Three of the seven genera dealt
with here are cold-water forms, temperate to subarctic in
distribution (Sebastes, Sebastolobus, and Helicolenus)
and the other four genera are tropical to subtropical.
Trachyscorpia will probably be shown to occur only in
the deep coastal waters of the southeast Pacific. The
single known specimen was taken from 580 to 600 m
(Chirichigno 1974) and listed as Trachyscorpia sp.

Over much of the California Current (CalCOFI) area,
the genera that commonly cooccur are Sebastes and
Sebastolobus. Although adults of Sebastolobus have
been taken as far south as Cape San Lucas, Sebastolobus
larvae are seldom taken south of California (Moser 1974).
Larvae of some species of Sebastes (e.g., S. macdonaldi)
are distributed primarily off Baja California (Moser
1972), but larvae of most species of Sebastes have a more
northern distribution.

In the area covered on EASTROPAC and STOR
cruises, between lat. 20°N and lat. 20°S, the larvae com-
monly taken are those of Pontinus and Scorpaenodes,
with larvae of Scorpaena collected occasionally and lar-
vae of Ectreposebastes found in equatorial waters as far
offshore as long. 126°W. To the south of the EAS-
TROPAC area, three temperate genera (Helicolenus,
Sebastes, and Trachyscorpia) occur. Larvae of Scorpaena
guttata are occasionally taken off southern California
and occur along Baja California south to Magdalena
Bay, hence cooccur commonly with Sebastes larvae. Lar-

Table 4- Length (rm) at hatching, notochord flexion, transformation into pelagic
juveniles, and known lengths of pelagic juveniles of scorpaenid
fishes included in this guide.

Length at

Length of

hatching

Length at

Length at

pelagic

Species

or birth

flexion

transformation

juveniles

Sebastes paucispinis

4.6

7.2- 9.7

15

15-34

4.0-5.0

7.7- 9.0

5.4

8.0-10.0

27-30

30-63

levis

5.0

7.6-10.4

19

19-58

cortezi

4.1

7.0- 8.3

ca 17

Gulf Type A

4.2

7.0- 7.6

-

capensis
marinus

3.8

6.2- 7.0

-

6.7-7.2

8.5-11.8

ca 24

viviparus

5.4-5.8

7.3-10.6

fasciatus

5.8

8.5-10.0

-

"

Sebastolobus alascanus

2.6

6.0- 7.3

14-20

20-27

altivelis

2.6

6.0- 7.3

Helicolenus dactylopterus

<2.8
1.8

6.0- 7.9
4.0- 5.4

11-14

-

Scorpaenodes xyris

Scorpaena guttata
type A

1.9-2.0
<2.2

4.5- 5.7

4.0- 5.5

-

<2.3

4.1- 4.6

ca 15

15-27

Type B

-

<5.0- 5.5

ca 10

Ectreposebastes imus

<2.8

5.5

ca 28

vae of Pontinus and Scorpaenodes have only oc-
casionally been taken off Baja California and as far north
as the vicinity of Cedros Island. The larvae of four genera
may cooccur in the Gulf of California — Scorpaena, Scor-
paenodes, Pontinus, and Sebastes. Larvae of Sebastes
and Scorpaena are taken commonly in the upper and
middle Gulf, whereas larvae of Pontinus and Scor-
paenodes are taken in the middle and lower Gulf;
however, there is a broad area of overlap of the four
genera in the Gulf.

Larvae of scorpaenids are commonly taken at con-
siderable distance from shore. In the CalCOFI area, lar-
vae are taken as far offshore as 500 km, and on EAS-
TROPAC cruises even further offshore. Scorpaenid larvae
that are taken at considerable distances offshore often
belong to species that have an extended pelagic juvenile
stage. A prime example is Sebastolobus altivelis, which
spend about 20 mo in the pelagic environment from
spawning to settling and attain a length of up to 56 mm
as pelagic juveniles (Moser 1974). Another example is
Pontinus Type A, the common form in the EAS-
TROPAC area. The larvae grow to about 15 mm before
transforming to pelagic juveniles, which then attain
lengths up to 27.4 mm. There apparently is a large size
range over which juveniles of Pontinus Type A become
demersal, since bottom-caught specimens as small as
17.2 mm are present in collections. This may be related
to the extensive offshore distribution observed for this
scorpaenid, with nearshore individuals settling out first.

KEYS TO LARVAE OF EASTERN PACIFIC
SCORPAENID GENERA

The most obvious feature of the larval period of marine
fishes is change. Changes in form and pigmentation from
hatching to transformation into the juvenile stage range
from substantial to spectacular. Because of this it is im-
possible to write a dichotomous key that would allow
identification of a species or genus of scorpaenid during
all phases of its early life history. To overcome this we
have constructed two keys, one for larvae which have not
undergone notochord flexion and have not formed me-
dian fins and the other for larvae which have completed
notochord flexion. The characters used in these keys are
summarized in the preceding section and discussed in
detail in the generic and species accounts. Since larvae of
the eastern Pacific species of Helicolenus and Trachy-
scorpia were not available to us we have omitted these
genera from the keys.

Key to Early Preflexion Larvae of
Eastern Pacific Scorpaenid Genera'

dorsal and ventral margins, sometimes ex-
panded to a solid band Sebastolobus

lb. Melanophores on tail along ventral mid-
line only, or along dorsal and ventral mid-
line, but each series contains a number of
discrete melanophores 2

2a. Series of both dorsal and ventral margin
melanophores developed on tail ....
Sebastes (some spp.)

2b. Only ventral margin melanophores series

on tail 3

3a. Melanistic shield of pigment covering dor-
solateral surface of gut 4

3b. No melanistic shield covering dorsolateral
surface of gut; pigment restricted to deeply
embedded blotch 5

4a. Larvae 2.0 to 3.0 mm at hatching, 24 myo-
meres, voluminous finfold Scorpaena

4b. Larvae 4.0 to 6.0 mm at hatching, 26 or 27

myomeres, moderate finfold . . .Sebastes (some spp.)

5a. Melanophores on pectoral fins restricted to

distal margin of fin Scorpaenodes

5b. Melanophores on pectoral fins distributed

over entire blade of fin 6

6a. Depth of pectoral fin base 18 to 22% of

body length Ectreposebastes

6b. Depth of pectoral fin base 15 to 16% of

body length Pontinus

Key to Postflexion Larvae of
Eastern Pacific Scorpaenid Genera'

la. Parietal ridge bifurcate posteriorly, with

posterior (nuchal) spine large 2

lb. Parietal ridge ending in single spine (pari-
etal) or if two spines are present the second
(nuchal) is small and positioned basally ... 3

2a. Myomeres, 29-31; dorsal spines, XTV-

XVn Sebastolobus

2b. Myomeres, 24 or 25; dorsal spines, XIII

Scorpaenodes

3a. Myomeres, 26 or 27; dorsal spines XHI . . . Sebastes

3b. Myomeres, 24; dorsal spines, XII 4

4a. Depth of pectoral fin base, 19 to 22% of

body length Ectreposebastes

4b. Depth of pectoral fin base 8 to 18% of body

length 5

5a. Pectoral fins wing-shaped Pontinus

5b. Pectoral fins with rounded posterior mar-
gin Scorpaena

DESCRIPTION OF LIFE HISTORY SERIES

la. Melanophores on tail portion of body re-
stricted to two large opposing blotches on

'Not included is Helicolenus from the southeastern Pacific which prob-
ably would key to couplet 4a based on the Atlantic species and Trachy-
scorpia which might key to la.

Of the 19 life history series included in this guide, 11
have not appeared previously in the literature or have
appeared as fragmentary accounts. These are Sebastes
jordani, S. levis, S. cortezi. S. sp. (Gulf of California,

'Trachyscorpia probably would key to couplet la and Helicolenus to 5b.

Type A), S. capensis{?), Scorpaenodes xyris, Scorpaena
guttata, Scorpaena Type A, Pontinus Type A, Pontinus
Type B, and Ectreposebastes imus.

For each genus there is a summary of the literature and
a section in which the distinguishing features of the lar-
vae and early juveniles are summarized and compared
with related genera. Our format for the species descrip-
tions consists of a literature summary, a section entitled
distinguishing features which contains the body of the
description, and a description on distribution. Where
there was adequate material, new descriptions are given
in more detail than are those which have appeared in the
literature previously. Morphometric tables and illus-
trations of developmental stages are included for all
species. Meristic tables were prepared for only eight of
the series: Sebastes paucispinis, S. macdonaldi, S. jor-
dani, Sebastolobus altivelis, S. alascanus, Scorpaenodes
xyris, Pontinus Type A, and Pontinus Type B. For the
others, there were either too few specimens to sacrifice
for staining or most of the specimens were leached of cal-
cium and would not stain. Where there is no meristic
table, data on fin counts are given in the distinguishing
features section.

Sebastes Cuvier

Literature. — Larvae of the genus Sebastes are the
best known of all scorpaenid genera although descrip-
tions of only a few species have been published. Plank-
tonic larval stages of the North Atlantic redfish, Sebastes
marinus, were first described and illustrated by Collett
(1880). This was followed by other descriptions and il-
lustrations of intraovarian stages (Ryder 1886; William-
son 1911; Goodchild 1924; Liiling 1951; Templeman and
Sandeman 1959) and planktonic stages (Dannevig 1919;
Jensen 1922; Bigelow and Welsh 1925; Einarsson 1960;
Taning 1961) of that species. The taxonomy of North At-
lantic Sebastes has long been in a state of confusion and
it is likely that some of the early larval descriptions are
referable to species other than S. marinus. Templeman
and Sandeman (1959) reviewed these papers and at-
tempted to correct the mistakes of the earlier workers. In
the papers by Einarsson (1960) and Taning (1961), the
larvae of the smaller North Atlantic species, Sebastes
viviparus, were described, illustrated, and compared
with those of S. marinus.

Eigenmann (1892) was the first to describe the in-
traovarian larvae of a species of Sebastes from the North
Pacific, S. rubrivinctus, and this was followed by de-
scriptions and illustrations of intraovarian larvae of other
northeastern Pacific species (Wales 1952; Morris 1956;
DeLacy et al. 1964; Ahlstrom 1965; Moser 1967a, 1972;
Waldron 1968; Efremenko and Lisovenko 1970;
Westrheim 1975). Complete planktonic larval series of
only 2 of the 65 northeastern Pacific species of Sebastes,
S. paucispinis and S. macdonaldi, have been described
and illustrated (Moser 1967a, 1972).

Larvae of the less speciose sebastine fauna off Japan
have been described by a number of investigators. Fujita

(1957, 1958) raised two species, S. pachycephalus nig-
ricans and S. oblongus, in aquaria up to the stage of fin
formation. Shiokawa and Tsukahara (1961) raised S.
pachycephalus pachycephalus from egg to juvenile, the
only time that a species of Sebastes has been raised
through transformation. Planktonic larval series of two
other Japanese species, S. hubbsi and S. inermis, have
been described by Uchida et al. (1958) and Harada
(1962), respectively. Takai and Fukunaga (1971) describ-
ed the larval development of S. longispinis, and recently
Sasaki (1974) has described preextrusion larvae of S.
schlegeli, S. steindachneri, and S. taczanowskii.

Distinguishing features. — Despite the large species
complement and resulting variety in larval form and pig-
mentation, there is a group of characters that allow
Sebastes larvae to be distinguished from those of the
other eastern Pacific genera.

In the earliest stages of larval development, there is a
marked dichotomy in size between Sebastes and the
other genera. At birth, Sebastes larvae are 3.8 to 7.5 mm
long, have utilized all or most of their yolk, and have
well-developed eyes and jaws. Where it is known, em-
bryonic development of the other eastern Pacific genera
occurs in floating egg masses and, at hatching, the lar-
vae are 2.0 to 3.0 mm long, have large elliptical yolk sacs,
poorly developed eyes and jaws, and a voluminous fin-
fold that has an inflated appearance. Development is
more advanced in relation to body length in these genera
compared with Sebastes. For example, the size at initia-
tion of notochord flexion ranges from 6.2 to 8.5 mm
(mean of 7.5 ± 0.72 SD) for 10 species of Sebastes studied
and at completion of flexion ranges from 7.0 to 11.8 mm
(mean of 9.5 ± 1.39). Comparative sizes for the other
genera are 4.0 to 6.0 mm (mean of 4.8 ± 0.97) at initia-
tion of flexion and 4.5 to 7.9 mm (mean of 6.1 ± 1.32) at
completion of flexion.

The extent and pattern of melanistic pigmentation is
useful in distinguishing Sebastes larvae from those of
other eastern Pacific genera. Intraovarian embryos
develop a shield of melanophores over the dorsolateral
surface of the gut and a series of melanophores along the
ventral midline of the tail. Almost half of the known
species also develop a series of melanophores along the
dorsal midline of the tail before hatching. Melanophores
may be added to the tail series after hatching, but the
length, anteroposterior position, and number of constit-
uent melanophores of these rows are usually stabilized
at the completion of yolk absorption (Table 5). This com-
bination of characters may be diagnostic for a particular
species but, usually, species identification requires sup-
plementary characters such as the pattern of melano-
phores on the pectoral fins and the presence or absence
of melanophores on the lower jaw, brain lobes, and
nape.

Larvae of Sebastolobus can be separated from
Sebastes on the basis of the tail pigment described
above. Early Sebastolobus larvae have two large
melanistic blotches midway along the tail, which disap-

Table 5. Mean number and range of midline melanophores on tail

portion of body of 33 species of Sebastes. All larvae were

collected from identified pregnant females.

Species of
Sebastes

Number of

specimens

examined

Ventral row

Dorsal row

X

range

X

range

melanostomus

27

8

4-11

0

0

macdonaldi

60

8

6-14

0

0

paucispinis

20

9

5-14

0

0

flavidus

20

10

8-12

0

0

ensifer

20

11

8-17

0

0

miniatus

40

11

8-18

0

0

eos

20

12

8-14

0

0

wilsoni

20

12

9-15

0

0

rosaceus

20

12

8-14

0

0

chlorostictus

20

13

11-14

C

0

rosenblatti

14

14

11-16

0

0

constellatus

20

14

11-17

0

0

serranoides

3

14

12-15

0

0

elonqatus

20

17

12-19

0

0

diploproa

20

17

13-20

0

0

levis

20

17

13-22

0

0

brevispinis

20

18

15-23

0

0

serriceps

27

19

15-24

0

0

goodei

11

20

18-24

0

0

umbrosus

20

22

17-29

0

0

rufus

20

9

5-13

1

0-3

oval is

20

12

10-15

15

12-18

marinus

18

18

11-24

13

8-21

jordani

18

24.

21-27

19

13-28

hopkinsi

20

26

22-31

13

9-18

dallii

20

29

25-34

12

8-17

caurinus

20

42

35-58

14

9-18

saxicola

20

43

34-55

10

7-16

gilli

20

46

32-69

8

3-13

semicinctus

20

50

40-63

20

15-23

auriculatus

20

52

39-63

25

15-31

carnatus

20

55

49-63

16

11-23

maliqer

20

63

54-69

13

8-19

pear in larvae 5.0 to 6.5 mm long. The warm-water
genera, Ectreposebastes, Pontinus, Scorpaena, and Scor-
paenodes, develop a row of about a dozen melanophores
on the ventral midline of the tail in very small larvae but
these are lost before the completion of notochord flexion.
The dorsal midline row that forms on some species of
Sebastes never appears on the larvae of these genera.

Late-stage larvae of Sebastes can be differentiated
from those of other eastern Pacific genera on the basis of
meristic characters. Eastern Pacific species, with a single
exception, normally have 26 vertebrae and 13 spinous
dorsal rays. The exception, S. polyspinis, has 28
vertebrae and 14 spines. Sebastolobus has higher counts
(29 to 31 vertebrae and 15 to 17 dorsal spines) and the
warm-water genera have lower counts (24 vertebrae and

12 dorsal spines). The only departure from this
generalization is Scorpaenodes which, like Sebastes, has

13 dorsal spines.

Of the 69 known eastern Pacific species, complete

developmental series, from newborn larvae to juvenile,
are described only for S. paucispinis (Moser 1967a) and
S. macdonaldi (Moser 1972), however, intraovarian and
newborn larvae have been described and/or illustrated
for a total of 39 eastern Pacific species. This paper con-
tains illustrated descriptions of S. paucispinis, S. mac-
donaldi, S. jordani, S. levis, and two larval forms from
the Gulf of California. Also presented are illustrations of
pre- or postextrusion larvae of 23 other eastern Pacific
species (Figs. 1-4). For comparative purposes, brief illus-
trated descriptions of Sebastes larvae from other regions
are also given. These include eight Japanese species, a
larval form from off Chile (probably S. capensis), and
three species from the North Atlantic. Research on the
larvae of the latter group is problematic because of the
taxonomic confusion that surrounds the adults. Adults
of the widely distributed redfish, S. marinus, are readily
distinguished from those of S. viuiparus, the diminutive
shallow-water form of the northern European coast. The
problem has been whether the deep-water "mentella"
form, which cooccurs geographically with S. marinus, is
specifically distinct from S. marinus and whether the
"American" form is a distinct species. The work of Bar-
sukov (1968, 1972) and Barsukov and Zakharov (1972)
indicates that S. marinus, S. mentella, and the "Ameri-
can" form, S. fasciatus, are distinct species, making a
total of at least four species in the North Atlantic. Esch-
meyer's systematic review of North Atlantic redfishes (in
progress, cited in Hallacher 1974) confirms this.

Sebastes pauciapinia (Ayres), Figure 5

Literature. — Newly hatched larvae of the bocaccio,
S. paucispinis, were illustrated by Morris ( 1956) and lar-
val stages up to 12 mm were illustrated in Ahlstrom
(1965). A complete description of the development of em-
bryos, larvae, and juveniles was given by Moser (1967a).

Distinguishing features. — Bocaccio larvae hatch and
are extruded at a length of 4.0 to 5.0 mm. Beginning at
about 15 mm they undergo certain changes (e.g., diminu-
tion of parietal and preopercular spines and develop-
ment of dorsal pigment saddles) that mark the begin-
ning of the pelagic juvenile stage. Transformation into
demersal juveniles occurs at 30 to 35 mm in length.

The larvae are comparatively slender. Body depth at
the base of the pectoral fin averaged 20^^ of the body
length before notochord flexion, 23% during flexion, and
30% in larvae which have completed notochord flexion
(Table 6). They develop the complement of head spines
typical for all species of Sebastes (Moser 1967a).

The most distinct morphological features of bocaccio
larvae are the elongate paired fins. The pectorals begin to
elongate and become aliform soon after hatching. They
reach a maximum relative length of about 37% of the body
length in larvae about 13 mm long. They gradually
diminish in relative length during the remainder of the
larval and pelagic juvenile stages and are 25% of the body
length in late pelagic juveniles. Ossification of the rays
begins in 6-mm larvae and the unusually small adult

Figure I.— Larvae of Sebaates (subgenus Sebattomua) removed from pregnant females and cultured to
the point of yolk exhaustion. A. S. constellatu*. 4.2 mm; B. S. rotaceiu, 3.9 mm; C. S. eiuifer, 4.2 mm; D.
S. umbroaus, 4.2 mm; E. S. chloroiticttu, 4.2 mm; F. S. roaenblatti, 4.6 mm; G. 5. eos, 4.4 mm.

10

Figure 2.— Larvae otSebastes at stage of yolk exhaustion. A. S. melanoatomus, 4.4 mm; B. S. miniatus,
4.3 mm; C. S. elongatus, 4.9 mm; D. S. ■errtcep*, 4.8 mm; E. S. goodei, 5.8 mm.

11

B

Figure 3.— Larvae of Sebastes at stage of yolk exhaustion. A. S. rufus, 5.3 mm; B. 5. ovalis, 4.5 mm; C.
5. hopkiiui, 4.7 mm; D. S. doUtt, 4.9 mm; E. S. aemicinctus, 5.5 mm; F. S. carnatua, 4.6 mm.

12

B

Figure 4.— Larvae of Sebastet at stage of yolk exhaustion. A. S. pinniger, 4.0 mm, redrawn from Waldron (1968);
B. S. aaxicola, 4.7 nun; C. S. auriculatiu, 5.2 nun; D. S. cauriniu, 5.5 mm; E. S. gilli, 5.2 mm. Specimens C and D
supplied by C. R. Hitz.

13

Figure 5. — Developmental series o( Sebaatee paucUpinU. A. 4.7-mm larva; B. 6.1-mm larva; C. 8.7-min
larva; D. H.O-mm larva; E. 24.2-mm pelagic juvenile.

14

Measurements (i«in) of Sebastes pauctspinis larvae. {Specimens betnrem
dashed lines are undergoing notochord flexion. )

standard
length

Snout-
distance

Head
length

Snout
length

Eye
diameter

eody
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin
distance

4.6

1.7

1.1

0.3

0.4

O.B

0.5

0.4

-

i.B

1.9

1.2

0.3

0.4

0.9

0.6

0.4

-

5.1

2.1

1.4

0.4

0.4

I.O

o.e

0.5

0.1

-

s.e

2.5

1.7

0.4

0.5

1-2

1.2

0,5

0.2

-

6.6

2.9

1.9

0.6

0.7

1.4

1.4

0.6 J

7.2

3.2

2.1

0.6

0.6

1.7

1.8

0.7

7.5

3.0

2.0

0.6

0.6

1.6

1.9

0.7

a.2

3.5

2.4

0.7

0.8

1.8

2.3

0.7

4.9

8.8

4.2

2.7

0.9

0.8

2.1

2.4

0,8

5.2

9.7

4.8

3.1

1.0

0.9

2.3

3.0

0,8

6.0

10.3

5.3

3.8

1.2

1.2

2.9

3.7

0.8

6.3

10.5

5.7

3.9

1.3

1.2

3.1

3.6

0.8

6.9

1!.0

7.0

4.5

1.4

1.4

3.5

4.4

1.0

7.9

12.7

7.1

4.6

1.5

1.5

3.7

4,6

1,0

8.1

13.7

8.2

5.0

1.4

4.1

5.1

1.0

9.2

14.2

8.6

5.5

1.8

1.7

4.2

4.8

1,0

9.2

15.0

9.1

5.5

1.7

1.8

4.6

5.4

1,1

9.7

24.2

15.2

8.8

2.9

2.6

6.9

8.3

1.3

29.4

19.0

11.3

2.9

3.0

8.1

7.4

1.7

•32.0

19.2

11.7

3.4

3.0

8.8

8.3

1,7

7.8

•34.4

20.6

11.5

3.8

3.1

9.4

8.8

2,0

7.5

22.3

as far north as Kodiak Island, although the principal dis-
tribution of the species is off California; they have been
reported offshore to depths of 320 m (Miller and Lea

■Transforming specimens.

complement of pectoral rays is present in 9-mm larvae
(Table 7). The pelvic fins begin to develop in 5-mm lar-
vae and, like the pectorals, reach a maximum relative
length of 37*^0 of the body length in 13-mm larvae.

Bocaccio larvae may be readily identified by their pig-
ment pattern. The most striking melanistic pigment is
on the posterior margins of the pectoral and pelvic fins.
This pigment remains throughout the larval period and
is found in juveniles up to about 70 mm. Other patches of
melanophores develop above the brain, on the opercle,
and at the base of the caudal fin. The number of
melanophores in the ventral midline series is small com-
pared with other species of Sebastes. Their number rang-
ed from 6 to 14 with a mean of 9 for 20 larvae counted.

Distribution. — Adults of S. paucispinis have been
taken as far south as Punta Blanca, Baja California, and

1 {'

< 1 1 , < 1 ■■'- 1 1

' '

-

o /

c*ce

UtNOOCIMO

■

.

o

-

^

■^ ^yc SAN

64 FRMCISCO

■

-

^^

■

\

X y CONCEPTION

■

~

\

o JZ_ \

-

■

"^

\o o oO° <s\ L

■

_

\ °°^ V

\

-

\ °'^\

\ ' o \

\ *^J

J^\

\ ° "^

^ \

1.

.

\ /^A °

*Y\

s

-

\

\.

-

.^ ^

'^.

-

V

-

■

_ 1 i

1 1 L

1 1 1 1 1 1 1 I 1

- 2

i 1

Figure 6. — Stations at which larvae of SebaateB pcuicispinU were col-
lected by CalCOFI plankton surveys during 1953 and 1956. Solid
circles indicate stations where number of larvae exceeded mean num-
ber (6.7) for all positive stations. Area of frequent occupancy is out-
lined (see Ahlstrom 1961 for complete grid).

Table 7. Meristics from cleared and stained larvae of Sebastes paucispinis.

Principal

ProGurrent

Branchlo-

Pectoral

Gil!

Pelvic

caudal

caudal

stegal

fin

rakers

fin

Length

(mm)

fin rays

fin rays

rays

rayB

elements

(right
arch)

Anal
fin
rays

Dorsal
fin
rays

rays

Vertebrae

supe-
rior

Infe-
rior

supe-
rior

Infe-
rior

left

right

left

right

supe-
rior

Infe-
rior

upper
1 iinb

lower
limb

left

right

6.6

3

4

10

10

3

1-2

7.2

5

-

5

12

12

-

.

-

6

_

1-3

1-3

e.8

7

-

1

6

14

14

-

-

-

7

_

_

1-4

1-4

4

9.3

7

-

-

7

15

15

-

2

11

-

_

1-5

1-5

22

10.3

8

7

2

3

7

15

15

2

2

12

1-7

IV-9

1-5

1-5

23

11.0

8

4

4

7

15

15

2

4

13

II-9

XIV-14

1-5

1-5

26

11.8

8

6

7

7

15

15

2

4

14

II-9

XIII-14

1-5

1-5

26

12.0

8

7

8

7

15

15

2

5

15

II-9

XIII-14

1-5

1-5

26

13.4

8

8

8

7

15

15

2

5

15

II-9

XIII-15

1-5

1-5

26

14.2

8

11

11

7

15

15

2

6

17

rii-9

XIII-14

1-5

1-5

26

15

1972). Information from CalCOFI samples shows that
bocaccio larvae occur as far south as Punta San Hip-
polito, Baja California, and seaward to about 400 km
(Fig. 6). The distribution of bocaccio larvae is not known
north of California since the CalCOFI sampling pattern
usually is terminated at the California-Oregon border.
Bocaccio larvae are present in CalCOFI samples for an 8-
mo period from November to June with a peak abun-
dance in January and February. Seasonal changes in
abundance and size of larvae are shown in Table 8.

Table 8. Mean lengths (onn) of Sebasteg pauciaplnie larvae collected on crulaes
of the California Cooperative Oceanic Fisberiea Inveatigationa during 1953
and 1956. (standardized numbers of larvae are shown in parentheses.)

Table 9. Measurements (mm) of larvae and pelagic juveniles of Sebastes macdonaldi .
(Specimens between dashed lines are undergoing notochord flexionV)

JiS

FEB

HAS

*PH

MAT

jm

SOT

sec

1953

5.3(88)

5.2(221)

8.2(82)

10.2(1,7)

13.2(5)

12.2(10)

1,.0(7)

1..5(56)

1956

1..9(89)

5.14(1.3)

7.8(27)

8.1,(19)

15.7(12)

7.9(5)

-

li-5(37)

Sehastea macdonaldi (Eigenmann and Beeson),
Figure 7

Literature. — A description of the larvae and pelagic
and benthic juveniles of S. macdonaldi, the Mexican
rockfish or "coral cod," is presented in Moser (1972).

Distinguishing features. — Larvae of S. macdonaldi
are born at a length of 4.0 to 5.0 mm. The gradual trans-
formation into pelagic juveniles begins at about 15 mm.
The pelagic juvenile phase appears to be highly pro-
tracted, since the smallest demersal juveniles known are
about 60 mm in length.

Larvae of this species are relatively deep-bodied. Body
depth at the base of the pectoral fins averages 23% of the
body length before notochord flexion, 33% during
notochord flexion, and 34% in larvae which have com-
pleted notochord flexion (Table 9). The sequence of
development of the head spines is described in detail in
Moser (1972).

The pectoral fins are short and compact, although they
gradually increase in length during the larval period. Fin
length averages 8% of the body length before notochord
flexion, 13% during flexion, and 19% in larvae which
have completed flexion. The fin elongates markedly in
pelagic juveniles, where its average length is 31%i of the
body length. The sequence of ossification of the fins and
bony skeleton is given in Table 10.

The most distinctive pigment is on the paired fins. The
pectorals are covered solidly with melanophores in new-
born larvae. When rays develop the solid pigmentation is
restricted to the membranes between the rays but the
overall appearance is a dark compact fin with light
striations. The pelvic fins develop similar pigmentation.
When the paired fins become elongate in pelagic
juveniles, they retain their distinct dark pigmentation.
Early larvae have a group of melanophores on the nape
and an unusually low number of melanophores in the
ventral tail series. In a sample of 60 newborn larvae, the
number of ventral midline melanophores ranged from 6
to 14 with a median of 8. Patches of melanophores

standard
length

Snout-Anus
distance

Head
length

Snout
length

Eye
diameter

Body
depth

Pectoral
fin
length

Pelvic

fin
length

Snout-anal

fin

distance

4.5

1.8

I.l

0.38

0.37

0.09

0.27

4.7

1.8

1.2

0.30

0.45

1.0

0.40

5.0

1.8

1.2

0.30

0.46

1.0

0.33

-

5.2

Z.l

1.4

0.48

0.51

1.2

0.36

5.7

2.4

1.6

0.47

0.57

1.3

0.52

5.9

2.6

1.8

0.48

0.55

1.5

0.45

-

6.3

2.8

1.8

0.56

0.57

1.6

0.60

0.05

6.7

2.9

1.9

0.56

0.63

1.7

0.61

0.04

-

6.8

3.1

1.9

0.55

0.70

1.8

0.62

0.15

.

7.0

3.3

2.1

0.75

0.70

1.9

0.60

0.12

-

...lA....

__3.5

...?;?..

..QiZS..

...0.7.3...

..2.O..

....0.68.,

...Q.2Q.

.

7.7

3.8

2.7

0.82

0.S4

2.3

0.90

0.33

4.2

7.9

3.8

2.7

0.80

0.90

2.5

0.93

0.25

4.2

8.2

4.2

2.9

1.1

1.0

2.5

1.1

0.52

4.8

8.5

4.7

3.2

1.2

1.0

3.0

1,3

0.80

4.9

9.0

5.0

3.2
........

1 .1

1.0

2.8

1.2

..........

0.65

5.4
..........

..........

...-J.J.....

"""5:89'

9.5

5,2

3.6

1.3

1.2

3.1

1.5

1.1

5.6

9.7

5.4

3.5

1.2

1.2

3.2

1.6

1.1

5.8

10.0

6.2

3.7

1.2

1.3

3.5

1.8

1.2

6.4

10.3

6.3

3.8

1.4

1.3

3.5

1.7

1.2

6.5

11.2

6.8

3.8

1.4

1.3

4.0

2.1

1.5

7.0

12.0

7.7

4.5

1.4

1.5

4.2

2.4

1.8

7.7

15.0

9.4

5.8

1.7

1.9

5.2

3.5

3.1

9.7

15.4

9.3

5.8

1.7

1.8

5.5

4.0

2,9

9.6

15.9

10.0

6.5

2.1

2.0

5.2

3.6

2,8

10.1

•22.6

14.0

8.4

2,7

2.3

7.5

6.7

4,8

14.7

•27.8

18.6

10.6

3.1

2.8

8.9

8.8

6,2

19.3

•29.2

19.1

10.8

2.9

3.2

10.0

9.8

6,5

19.9

•32.7

20.7

11.8

3.2

3.6

10.1

9.7

-

20.8

*34.5

20.9

11.8

3.2

4.0

10.5

10.6

7.7

21.4

'38.8

25.0

12.8

3.2

4.0

11.7

11.2

8.3

25.6

'40.5
•44.4

26.8
30.2

13, 0
14.0

3.5
3.8

4.2

4.7

12.0
13.2

12.2
13.2

8.5
9,2

27.4
31,2

•Pelagic juvenile,

develop over the brain, on the opercle and preopercle.
Late in the larval period, melanophores form lateral to
the base of the dorsal fin. It is from these elongate
patches that the striking pigment saddles extend ven-
trad in pelagic juveniles.

Distribution. — This species has the most southerly
distribution of any species of Sebastes in the eastern
North Pacific. Adults have been taken as far north as
Pt. Sur, Calif., and as far south as Morgan Bank, near
Cape San Lucas, Baja California. Its abundance on the
banks off southern Baja California, and in certain areas
of the Gulf of California, suggests that the outer coast
populations and the Gulf populations may be continuous
(Moser 1972).

CalCOFI data show that larvae of this species are most
abundant off central Baja California. Larval abundance
declines sharply to the north of this region and larvae do
not occur north of Los Angeles (Fig. 8). Larval abun-
dance falls off less sharply to the south of Punta Eugenia;
however, the CalCOFI sampling pattern does not allow
exact delimitation of the southern range of S. mac-
donaldi larvae.

The larvae are present in CalCOFI samples over a 6-
mo period from January to June with a peak abundance
in March. Seasonal changes in abundance and size of lar-
vae are shown in Table 11. A few larvae of this species
were taken in the Gulf of California.

16

Figure 7.— Developmental series of Sebaatea macdonaldi. A. 4.S-mm larva; B. 6.3-mm larva; C. 9.0-min larva; D.
15.4-miii transforming specimen; E. 29.2-mm pelagic juvenile.

17

Table 10. Meristics from cleared and stained larvae of Sebastes macdonaldl.

Length

Principal
caudal
fin ray a

Procurrent
caudal
fin rays

Branchlo-
stegal
rays

Pectoral
fin
rays

Hypural
elements

G111
rakers
(right

arch)

Anal
fin

Dorsal
fin

Pelvic
fin
rays

Vertebrae

supe-
rior

infe-
rior

supe-
rior

infe-
rior

left

right

left

right

supe-
rior

inte-
rior

upper

linb

lower
Umb

rays

rays

left

right

6.3

.

2

2

2

.

.

_

.

.

_

6.7

-

-

-

-

3

3

-

-

-

-

-

-

-

-

6.8

-

-

-

-

3

-

-

-

-

4

-

_

-

-

7.0

3

-

-

6

6

-

-

-

5

-

-

-

7.1

«

-

-

6

6

-

-

-

6

-

-

-

7.2

3

-

-

8

-

-

-

6

-

-

-

-

7.7

6

-

-

11

11

-

-

9

_

-

I-O

1-0

5

7.9

1

-

-

9

10

-

-

9

_

-

3

8.2

8

2

2

17

17

-

-

1

12

III-4

VII-0

1-3

1-3

24

8.5

7

-

15

15

1

1

1

11

.

1-3

1-3

18

8.7

8

-

16

16

1

12

-

IV-0

1-3

1-3

18

9.2

8

2

19

18

2

3

14

-

VII-8

1-4

23

9.4

8

4

19

19

2

3

15

III-7

xin-13

1-5

1-5

24

9.7

8

3

19

19

2

3

16

in-7

XIII-13

1-5

1-5

24

10.0

8

5

19

19

2

5

17

in-7

)(III-13

1-5

1-5

26

10.3

8

5

19

20

2

-

-

in-7

xni-13

1-5

1-6

26

11.3

8

4

19

19

2

5

18

III-7

xin-13

1-5

26

12.0

8

8

19

19

2

5

18

III-7

XIII-13

1-5

1-5

26

15.0

8

10

11

7

19

19

3

-

-

III-7

xin-13

1-5

1-5

26

15.9

8

10

11

7

19

19

3

9

21

III-6

XIII-12

1-5

1-5

26

27.8

-

-

-

-

18

18

2

2

10

25

III-7

XIII-13

1-5

1-5

26

_L

Table 11. Mean lengths (mm) of Sebastes macdonaldl larvae coUected on cruises

of the California Cooperative Oceanic Fisheries Investigations

during 1953 and 1960. (Standardized numbers of larvae

are shown in parentheses.)

Figure 8. — Stations at which larvae of Sebastes macdonaldi were col-
lected during 4 yr (1953. 1%0. 1965. and 1966) of CalCOFI plankton
surveyB. Solid circles indicate stations where number of larvae ex-
ceeded mean number (4.8) for all positive stations. Area of Sequent
occupancy is outlined (see Ahlstrom 1%1 for complete grid).

JAN

FEB

HAP

APR

HAY

JUN

1953

4.3(45)

4.6 (109)

4.9 (632)

5.6 (163)

5.2 (22)

7. 8 (2)

1960

4.2 (3)

4. 8 (65)

5.1 (1042)

5.1 (248)

5.2 (48)

4.6 (15)

Sebastes jordani Gilbert, Figure 9

Literature. — Morris (1956) described briefly the new-
ly extruded larvae of S. jordani and illustrated a 6.8-nim
specimen.

Distinguishing features. — The larvae of the short-
belly rockfish, S. jordani, are larger at hatching than
other eastern Pacific species studied to date. Morris
(1956) illustrated a newly hatched S. jordani larva 6.8
mm in length, however, the mean for 20 measured
specimens was 5.4 mm. This is 0.75 mm larger than the
mean length for the next largest species listed by Morris.
Newly hatched larvae of most other eastern Pacific
species are between 3.8 and 5.0 mm. The 6.8-mm length
of Morris' illustrated specimen is above the upper range
of any eastern Pacific species known to us and is more
like the lengths of newly hatched larvae of the S. marinus
group of the Atlantic and some Japanese species. Other
similarities between the larvae of S. jordani and the S.
marinus group suggest that the two may be closely
related. Sebastes jordani larvae have an extended larval
period. Transformation into the pelagic juvenile stage oc-
curs at about 27 mm and specimens in the 28- to 30-mm

18

:•:■■::^/^;■:;^■;■; •"■v>*f^f<g|-;,i^^^%,^

Figure 9.— Developmental series of Sebcutes jordani. A. 7.2-mm larva; B. 10.0-mm larva; C. 15.5-min larva; D.

21.0-mm larva; E. 29.6-mm pelagic juvenile.

19

size range have the morphological and pigmentary
features of pelagic juveniles. This is also true for S.
marinus. Pelagic juveniles remain in the surface waters;
they appear to form schools soon after transformation
and are particularly susceptible to dip netting under
nightlights. Juveniles in the 30- to 50-mm size range are
commonly dip netted and a 62.8-mm specimen was
taken by this method (Table 12).

Larvae of S. jordani are the most slender of all
Sebastes yet described. Body depth averages 17% of the
body length before notochord flexion, 21% during flexion,
and 23.5% following flexion. Pelagic juveniles 28 to 68
mm long appear to be slightly more slender, with a mean
of 22%.

The rounded pectoral fins are the shortest of any
eastern Pacific rockfish larvae yet described. Pectoral fin
length averages 7% of the body length prior to notochord
flexion and 8% during flexion. This percentage increases

Table 12. Measurements (nw) of larvae of Sebastes jordani. (Specimens between
dashed lines are undergoing notochord fleidon.}

Standard
length

Snout-
anus
distance

Head
length

Snoot
length

Eye

diajneter

Body
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin
disunce

5.4

2.0

1.2

0.28

0.46

0.85

0.38

0.32

...

...

S.8

2.1

1.2

0.31

0.45

0.95

0.32

0.38

...

6.1

2.2

1.3

0.30

0.49

1.0

0.43

0.43

...

6.4

2.4

1.6

0.4«

0.60

1.2

0.45

0.45

...

7.2

2.6

1.6

0.45

0.61

1.2

0.48

0.46

7.7

2.8

1.8

0.55

0.65

1.4

...8i55...

...5.50.

e.o

3.3

2.1

0.72

0.74

1.6

0.62

0.60

0.07

4.4

8.6

3.7

2.2

0.78

1.8

0.64

0.63

0.08

4.8

9.0

3.7

2.7

0.94

0.87

1.8

0.70

0.70

O.IO

5.2

9.4

4.2

2.8

1.0

0.92

2.1

0.83

0.75

O.IS

5.2

10.0

4.2

3.0

1.1

0.88

2.2

1.0

0.82

0.18

5.8

10.5

4.8

3.3

1.2

1.0

2.4

1.2

1.0

0.35

6.4

11.6

5.3

3.9

1.4

1.1

2.5

1.4

1.0

O.M

6.9

12.8

6.2

4.1

1.6

1.2

3.1

1.7

1.1

0.50

7.6

13.4

7.2

4.8

1.7

1.3

3.2

2.0

1.1

1.0

8.8

15.6

8.0

5.0

1.7

1.7

4.1

2.5

1.2

1.4

9.2

21.0

11.3

6.7

2.2

2.1

5.2

3.9

1.7

2.2

12.2

22.8

11.7

7.4

2.6

2.1

5.3

4.1

1.6

2.5

13.2

23.5

12.0

7.5

2.6

2.2

5.6

4.6

1.8

2.7

13.8

24.5

12.8

8.2

2.9

2.2

5.4

4.7

1.7

3.0

15.0

25.5

13.8

8.1

2.7

2.3

6.3

4.8

1.8

2.9

15.0

26.4

13.8

8.3

2.8

2.4

6.3

5.0

1.9

3.3

15.7

27.0

14.0

8.5

2.8

2.7

6.4

5.3

1.9

3.4

16.4

•28.4

15.0

9.5

3.2

2.6

6.4

5.4

2.0

3.3

17.2

•29.6

15.7

9.4

3.2

2.6

6.9

5.8

2.2

3.7

16.1

•30.3

16.7

10.0

3.8

2.6

6.7

6.2

2.1

3.8

19.3

•40.7

22.0

12.8

4.6

3.1

8.5

2.7

5.4

25.8

•50.7

26.5

15.0

4.8

4.2

11.2

11.0

3.3

7.5

31.5

•62.8

33.0

17.4

5.2

4.6

13.5

13.5

4.2

9.2

38.3

•Pelagic juvenile.

gradually following notochord flexion from 11% im-
mediately after flexion to 20% in the largest larvae.
There is only a slight increase in relative pectoral fm
length during the pelagic juvenile stage, up to 22% in the
largest specimen available (62.8 mm). The small pec-
torals in S. jordani and the ontogenetic changes in
relative size agree closely with the condition in the lar-
vae of the S. marinus group. The sequence of ossification
of the pectorals and other fins is shown in Table 13.

A character which separates adults and juveniles of S.
jordani from all other known species of Sebastes is the
anterior placement of the anus. In his key Phillips (1957)
indicated that the distance between the anus and the
first anal spine in S. jordani is 1.1 to 1.5 orbit diameters,
whereas this distance is 0.1 to 0.8 orbit diameters in other
species. This wide space between the anus and the anal
fin origin is conspicuous in pelagic juveniles and is a
useful character in identifying all but the smallest lar-
vae. Snout-anus distance averages 36% of the body
length in the larvae prior to notochord flexion, 42% dur-
ing flexion, and 51% following flexion. In pelagic
juveniles it is 54%. Comparative mean percentages for
larvae of S. paucispinis are 41 before flexion, 45 during
flexion, and 59 following flexion. For larvae of 5. mac-
donaldi, the percentages are 42, 52, and 60.

The pigment pattern of early S. jordani larvae is
similar to that of the S. marinus group in the Atlantic.
Newborn larvae have the melanistic shield over the gut,
a group of melanophores above the brain, and a series of
melanophores along the dorsal and ventral midlines of
the tail. The ventral row begins at the 3rd to 5th postanal
myomere and extends to the 14th and 16th postanal
myomere. Typically, in the anterior two-thirds of the
series, the melanophores are positioned singly, but in the
jTOsterior two-thirds the melanophores are in two ir-
regular rows along the ventral midline. For 15 specimens
counted, the total number in the ventral row ranged from
21 to 27 with a mean of 23.7 ± 1.75 SD. The dorsal row is
considerably shorter, beginning at the 7th to 12th
postanal myomere and extending posteriad to the 14th to
16th myomere. The melanophores are bunched together
in an irregular row that contains a mean of 18.9 ± 4.64
melanophores (range of 13 to 28) for 18 larvae counted.

Table 13. Meristics from cleared and stained larvae of Sebastes jordani.

Length

(nm)

Principal
caudal
fin raya

Procurrent
caudal
fin rays

Branchlo-
stegal
rays

Pectoral
fin
rays

Hypural
elements

rakers
(right
arch)

Ana!
fin
rays

Dorsal
fin
rays

Pelvic
fin
rays

Vertebrae

supe-
rior

infe-
rior

supe-
rior

infe-
rior

left

right

left

right

supe-
rior

infe-
rior

upper
limb

lower
limb

left

right

9.4
10.0
10.5
11.2
12.5
12.8
22.0

8
8
8
8
8
8
8

0
0
2

0
3

5

10

1

1
0
2
3

4

5

12

10
13
16
16
19
19
21

10
13
16
16
19
19
21

2
2
2
2
2

3
3
2
2
2

0
1
2
2
4
4
10

11
12
15
15
18
19
26

III-6
III-11
III-11

XI!I-6

XIII-13

Xin-14

1-3
1-3
1-5
1-4
1-5

18
20
24
27
27
27
27

20

Development of pigment pattern in later larval stages
is also similar to that in the S. marinus group of the At-
lantic. The ventral series on the tail becomes embedded
and obscure; beginning at the anterior end of the series
this process gradually proceeds posteriad until, in 15-mm
larvae, only those melanophores posterior to the anal fin
are visible. With further development, these too are ob-
scured. The dorsal series follows an opposite course.
Melanophores are added anteriorly to form streaks on
either side of the developing dorsal fin. Melanophores are
also added posteriorly so that the dorsum is pigmented
from head to caudal fin. Melanophores begin to appear
on the side of the tail just anterior to the caudal fin in lar-
vae of about 15 mm in length. Also, the posterior mar-
gins of the hypural elements become outlined with em-
bedded pigment.

At transformation, which begins in specimens about 27
mm long, the upper half of the body above the lateral line
begins to be covered with fine melanophores. These
melanophores are more concentrated at the myosepta
and accent them. The streaks along the soft dorsal fin
and the streak above the caudal peduncle are darker
than on the more anterior region of the dorsum. In 40-
mm pelagic juveniles, the general body pigmentation
begins to extend below the lateral line, but even in later
juveniles and in adults, the ventral region of the body is
paler than the dorsal half. The pectoral and pelvic fins
develop no melanistic pigmentation in larvae or pelagic
juveniles.

Distribution.— According to Miller and Lea (1972), S.
jordani ranges from British Columbia south to Cape Col-
nett, Baja California (about lat. 31°N). A 45.2-mm
juvenile (SIO 71-120) dip netted off west San Benito Is-
land (lat. 28°18.0'N, long. 115=34.25^) extends the
range of this species about 270 km south. The dis-
tribution of S. jordani larvae in the CalCOFI sampling
area during 1966 is shown in Figure 10. The sampling
area obviously stops short of the northward extent of
spawning, however, it shows the southern extent of lar-
val occurrence to be just south of the border with Mexico
(CalCOFI Line 100). Another obvious feature is the
coastal nature of S. jordani distribution. All larvae were
taken in the shoreward region of the sampling grid and

the largest individual catches were made on stations next
to the coast.

Frequency of occurrence of S. jordani is 7.7% off cen-
tral California and decreases to 5.7% off southern
California and 1.5% off northern Baja California (Table
14). Percentage of S. jordani larvae to total Sebastes lar-
vae was highest (20%) off southern California. Spawning
occurs over a short time period compared with other
species studied (Table 15); it extends from January to
April with a peak in February or March.

_L

Figure 10.— Stations at which larvae of Sebaates jordani were col-
lected by CalCOFI plankton Burveys during 1966. Solid circle* in-
dicate stations where number of larvae exceeded mean number (43.8)
for all positive stations. Area of ft'equent occupancy is outlined.

Table 14. Relative abundance and frequency of occurrence of Sebastes Jordani in CalCOFI collections for 1966.

CalCOFI regions and lines

Total
stations
occupied

Total

stations

containing

Sebastes

larvae

Percentage

of stations

containing

Sebastes

Total

stations

containing

S. jordani

Percentage
of stations
containing
S. jordani

Standardized

number of

S. jordani

larvae

Standardized

number of

Sebastes larvae

in hauls

containing

S. jordani

Percentage of

S. jordani

larvae to

total Sebastes

larvae

Central California
60-77

235

56

23.8

18

7.7

992

8,562

11.6

Southern California
80-93

511

253

51.5

29

5.7

1,355

6,822

19.9

Northern Baja California
97-107

459

122

26.6

7

1.5

40

692

5.8

Upper central Baja California
110-120

418

91

21.8

0

0

0

0

0

Lower central Baja California
123-137

325

36

11.1

0

0

0

0

0

21

Table 15. Stands
larvae tal

irdized numbers (A), mean length in mm with standard deviation (B), and size range in tm of Sebastes ,
ken on CalCOFI cruises of 1966. There was no cruise during March, and S. jordani were not ta'fceii
from May to December. Stations in central California were not occupied in February.

CalCOFI regions
and liiiee

January

February

April 1

A

B

C

A

B

C

A

B

C

Central California
60-77

781.9

4.9t0.59

4.0 - 7.0

-

205.2

7.5+3.31

5.1 - 24.5

Southern California
80-93

459.0

5.1 ±1.01

4.5 - 8.0

863.3

4.9 ±1.51

4.4 - 8.0

54.6

5.8 + 1.38

5.0 - 11.0

Northern Baja California
97-107

0

0

0

15.3

6.8 ±1.49

5.2 - 10.0

0

0

0

Sebastes levis (Eigenmann and Eigemnann),
Figure 11

Literature. — Newborn larvae of S. levis, the cow rock-
fish or cowcod, were illustrated in Moser (1967a).

Distinguishing features. — Larvae of S. levis are
about 5.0 mm long at birth; they achieve the relatively
large size of about 20 mm. A specimen 21.3 mm was just
beginning to develop the lateral pigment saddles typical
of pelagic juveniles, whereas transformation has
progressed far enough in a 19.1-mm specimen to classify it
as a pelagic juvenile. The pelagic juvenile phase appears
to be protracted since pelagic juveniles up to 58 mm were
taken in midwater trawls and the smallest demersal
juvenile found was 66.5 mm.

The large fanlike pectoral fins are diagnostic features
of cow rockfish larvae and attain the greatest size of any
species of Sebastes in the CalCOFI collections. Their
length increases from 9"^ of the body length in newborn
larvae to 46% at the completion of notochord flexion
(Table 16). Pectoral fin length in the few large larval
specimens is 41 to 45% of the body length. The large pec-
toral fins are characteristic of pelagic juveniles; 19- and
33-mm specimens had fin lengths 46 and 47% of the body
length. In larger pelagic juveniles the pectorals become
relatively smaller; fin length in a 47-mm specimen was
38% of the body length and in a 58-mm specimen it was
32%.

The melanistic pigmentation of cow rockfish larvae is
also an aid to identification. Newborn larvae have a mel-

Table 16. Measurements {mn) of Sebastes levis larvae. (Speciniens between dashed lines
are undergoing notocnord flexion.)

Standard

length

Snout-
anus
distance

Head
length

Snout
length

Eye

diameter

Body
depth

Pectoral
fin
length

Pectoral
fin base

depth

Pelwic
fin

length

Snout-anal

fin

distance

5.0

2.0

1.2

0.26

0.38

0.90

0.43

0.40

7.6

3.4

2.2

0.75

0.66

1.8

1.8

0.90

0.12

-

8.7

4.1

2.4

0.82

0.77

2.4

2.8

1.2

0.34

5.0

8.9

4.2

2.9

I.O

0.91

2.5

3.2

1.2

0.51

5.2

9.2

4.3

2.8

1.0

0.92

2.3

2.9

1-2

0.44

5.4

9.4

4.7

2.9

1.0

0.93

2.8

3.5

1.2

0.50

5.2

9.8

4.7

2.8

1.0

0.90

2.6

2.9

1.2

0.50

5.8

10.1

5.2

3.1

1.1

3.1

4.2

1.3

1-0

5.8

10.4

5.8

3.4

1.2

3-6...

4.8

1.3

1.2

6.2

11.3

6.4

4.3

1.2

3.7

5.0

1.3

1.8

6.8

12.3

7.0

4.2

1.3

4.2

5.1

1.5

2.2

7.4

21.3

13.5

7.2

2.1

7.1

9.6

2.4

5.1

13.8

•19.1

U.8

6.3

1.8

2.0

7.1

8.7

2.1

4.8

12.2

•32.8

20.9

10.8

3.0

2.8

11.2

15.4

3.5

9.2

21.9

•47.5

29.9

16.4

5.2

4.1

16.2

18.2

4.8

10.6

31.0

•58.4

36.6

20.1

6.7

5.0

20.3

18.9

5.4

13.0

38.5

•66.5

42.3

24.1

6.7

5.0

22.4

19.5

6.6

14.7

43.1

•Pelagic Juvenile.
"Benthic juvenile.

anistic shield over the gut, a patch of melanophores over
the brain, heavy pigmentation on the tip of the lower jaw,
and a moderate number of melanophores in the ventral
midline series (mean of 17 with a range of 13 to 22 for 20
specimens counted). The pectoral fin is covered with fine
melanophores that are concentrated in a band at the dis-
tal margin. With the rapid growth of the fin in larvae
between 6 and 7 mm, the melanophores on the major
part of the pectoral are lost but the marginal band
remains prominent. With further development, the mar-
ginal band widens so that in larvae of 11 mm the fin is
covered solidly with fine melanophores. This pigment is
retained in pelagic juveniles. When the larvae transform
into pelagic juveniles at about 20 mm, pigment saddles
begin to develop along the anterior region of the trunk.
Additional saddles develop posteriorly and with further
enlargement they become distinct bands. These are evi-
dent in pelagic juveniles as small as 33 mm and are
retained in demersal juveniles and adults.

Distribution. — Adult cow rockfish have a com-
paratively limited geographic range from central Califor-
nia to central Baja California with a principal dis-
tribution off southern California. The larvae are rare in
CalCOFI samples; only 30 (standardized number) were
found in collections from 1951, 1952, 1953, 1961, and
1969. Their geographic distribution suggests that, in
terms of reproduction, the cow rockfish is essentially a
species of the California Bight (Fig. 12). Cow rockfish lar-
vae were taken from January to June but in such low
numbers that months of maximum abundance could not
be determined.

Sebastes cortezi (Beebe and Tee-Van), Figure 13

Literature.— The literature contains no descriptions
of Sebastes larvae from the Gulf of California. The tax-
onomy of adult Sebastes in the Gulf is poorly known;
however, Chen (1975) has recently reviewed the species
of Sebastes from these waters. The larval series described
herein as S. cortezi was identified by raeristic characters
and transitional specimens provided by Chen.

Distinguishing features. — At birth, S. cortezi larvae,
like Sebastes larvae of the outer coast are 4.0 to 5.0 mm
long. The largest specimen in the series (17.1 mm) was
beginning to transform.

The most notable morphological features of S. cortezi
larvae are the short pectoral fins which have a slight dor-
sad orientation for most of the larval period, and the

22

Figure 11. — Developmental series otSebaateB levU. A. S.O-min larva; B. 7.2-mm larva; C. 10.4-
mm larva; D. 19.1-mm transforming specimen; E. 32.8-mm pelagic juvenile.

23

1 « ' ' ' ' 1 ' ' '

' 1

T^

•

r cft«

-

—

■

-

■

y^ ^^^ SA"

-

<c

-

- \

\^ POINT

-

-

\ \/

V.

-

■

\ ^>n*

)

-

\ •

\v

^

....

_

\

\n

, \

^

.

\ 1

\

>X'

K

-

\

1

\

V

-

\^

\

1

k.

\

-

\

^

i

4

-

^

s^

\

-

\>

-

■

-

-

1 1

1

Figure 12. — Stations at which larvae of Sebastes tevis were taken
during 4 yr (1951, 1955, 1968, and 1969) of CalCOFI plankton surveys.
Area of total grid is outlined.

length of parietal spines. Pectoral fin length is 7 to 9% of
the body length in larvae up to the beginning of
notochord flexion, 9 to 12% during notochord flexion, and
has increased to a maximum of 21% following notochord
flexion. It is 23% in the transforming specimen (Table
17). A morphological feature which helps to distinguish
S. cortezi larvae from Type A Gulf larvae is the length of
the parietal spines, the spines being relatively shorter in

Table \7. Measurements (nm) of larvae of Sebastes cortezi- (Specimens between aasned lines
are undergoing notocnord flexion.)

SUndard
lengtli

Snout-
anus
distance

Head
length

Snout
length

Eye
diameter

Body
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin
distance

4.)

1.7

1.0

0.25

0.34

0.75

0-37

0.28

...

4.5

1.8

1.2

0.30

0.38

0.95

0-30

0.40

4.;

1.9

1.2

0.37

0.47

1.2

0.37

0.38

5.1

!.2

1.5

0.43

0.49

1.3

...

5.7

2.5

1.7

0.48

0.54

1.3

0.41

0.57

6.1

2.6

1.7

0.52

0.55

1.4

0.57

0.60

6.4

2.9

1.9

0.63

0.60

1.6

0.60

0.64

0.06

7.0

3.5

2.1

0.68

0.78

0.65

0.82

0.07

4.2

7.2

3.7

2.2

0.78

0.85

2.0

0.73

0.90

0.10

4.2

7.8

3.8

2.5

0.75

0.86

2.1

0.77

0.91

0.14

4.5

8.0

4.0

2.7

0.75

-0.94

2.2

0.93

0.94

0.15

4-7

8.3

4.8

2.6

0.90

2.3

1.0

0.92

0.15

4.9

8.8

4.5

2.6

0.90

1.0

2.5

1.0

1.0

^ 0".16"

5.2

11.8

7.5

4.4

1.4

1.4

4.1

2.3

1.4

1.5

7.8

12.5

7.8

4.7

1,5

t-4

4.2

2.6

1.2

1.5

8.1

13.5

8.9

5.5

1-4

4.3

2.6

1.3

1-6

9.3

•17.1

10.3

6.3

1.9

1.8

5-6

3.9

1.6

2-5

10.8

S. cortezi. In larvae between 8.0 and 9.0 mm, when the
spines reach their greatest length relative to the head
length, those of S. cortezi are 21 to 22% of the head length
and those of Type A are 25 to 34%.

Melanistic pigmentation is more useful than mor-
phology in distinguishing S. cortezi larvae from other
rockfish larvae. At birth the larvae have the usual pig-
ment shield over the gut and a series of melanophores
along the ventral midline of the tail (mean of 15 with a
range of 10 to 21 melanophores for 15 larvae counted). A
large melanophore is located in the position of the future
caudal fin and remains there throughout the larval
period. The pectoral fins have an unusual pigment pat-
tern; there is a melanistic patch in the proximal ventral
sector of the fin and the dorsal and distal portions are
clear. With further development, the entire proximal
region becomes pigmented but the distal clear zone
widens. The medial surface of the fin base is covered with
large melanophores. The pelvic fins are pigmented in lar-
vae 11.8 mm and larger.

Other pigment develops above the brain and, late in
the larval period, on other regions of the head. In larvae
about 8.0 mm long a short series of five to six pairs of
melanophores develops along the dorsal midline of the
tail. With further development this series lengthens, and
extends along the entire dorsum in larvae 12.5 mm and
longer. The ventral series of melanophores becomes
restricted to the caudal peduncle after the anal fin is
developed (about 8.0 mm). The 11.8-mm larva has a
large patch of melanophores on the lateral surface of the
caudal peduncle. In later stages, melanophores extend
anteriad along the lateral line from this patch and, in the
17.1-mm specimen, the entire trunk above the lateral
line is becoming pigmented.

Distribution.— Eight CalCOFI cruises into the Gulf of
California in 1956 and 1957 and a U.N. Food and Agricul-
ture Organization (FAO) cruise in March of 1972 provide
information on the distribution and abundance of S. cor-
tezi larvae. Like the adults (Chen 1975), the larvae occur
in the western side of the Gulf and range latitudinally
from off Point San Fermin south to about Carmen Is-
land (Fig. 14). Most of the larvae were taken just north
and just south of the large islands, Tiburon and Angel de
la Guarda, in the upper Gulf. Although the results of the
various cruises are not strictly comparable because they
differed in area covered and in the number of stations oc-
cupied, Table 18 gives some indication of the seasonality
of spawning. Larvae are present during the cold-water
months and appear to have a maximum abundance in
March.

Table 18. Mean length (nin) with total standardized number of larvae of Sebastes
cortezi and S. Type A in p1anl(ton collections from the Gulf of California.

•Transfonning specimen.

Species

February

March

April

June

x length

Standard

number

ii length

Standard
number

» length

Standard
number

Standard

number

S. cortezi
S. Type A

4.3

5.2

31
177

7.2
6.5

121

75

6.2
6.6

23
25

0

0

24

Figure 13.— Developmental series otSebaatea cortezi. A. 4.5-mm larva; B. 6.8-mm larva; C. 8.8-mm larva; D.
ll.S-mm larva; E. 17.1-mm pelagic juvenile.

25

Table 19. Measurements (mn} of larvae of Sebastes sp, [Gulf of California Type A],
(Specimens between dashed lines are undergoing notochord flexion.)

Figure 14. — Stations in the Gulf of
California at which larvae of
Sebastes cortezi were talien on
CalCOFI cruises during 1956 and
1957.

Sebaates sp.— Gulf of California Type A, Figure 15
Literature. — See previous species account.

Distinguishing features. — At birth, Type A larvae
are 4.0 to 5.0 mm long. The maximum size obtained
before transformation is not known since the largest lar-
va in the collection is 10.8 mm.

Type A larvae, like those of S. cortezi, have short pec-
toral fins. Pectoral fin length is 6 to 9% of the body length
before onset of notochord flexion, and thereafter in-
creases to 20% in the longest larva (Table 19). At com-
parable sizes, the parietal spines are longer in Type A
than in S. cortezi.

Type A larvae have a distinctive pattern of
melanophores. As in S. cortezi, newborn larvae of Type A
have a shield of pigment over the gut and a series of
melanophores along the ventral midline of the tail (mean
of 16 with a range of 12 to 21 for 13 larvae). Unlike S. cor-
tezi larvae, those of Type A have a blotch of pigment at
the symphysis of the lower jaw, that persists in larvae up
to about 8 mm. The pectoral fins have fine melanophores
distributed over the entire blade of the fin and have the
medial surface of the fin base covered with large
melanophores. At about 8 mm, pigment appears above
the brain. No pigment forms on the dorsal surface of the
tail as in S. cortezi. The pigment pattern is similar to
that of the subgenus Sebastomus from outer coastal

Standard

length

Snout-

anus

distance

Head
length

Snout
length

Eye

diameter

Body
depth

Pectoral
fin
length

Pectoral
fin base

depth

Pelvic

fin

length

Snout-anal

fin

distance

4.2

1.5

0,84

0.23

0.33

0.7S

0.33

0.22

4,7

1.8

1,00

0.28

0.35

0.80

0.30

0.32

5.4

2.2

1.3

0.37

0.45

1.1

0.47

0.43

5.8

2-4

1.5

0.41

0.48

1.2

0.54

0.55

6.1

2.6

1.7

0.48

0.50

1.4

0.61

0.58

...

—

6.4

2.6

1.7

0.50

0.55

1.4

0.63 ,

0.60

...

7.0

3.6

2.4

0.90

0.73

2.1

0.90

0.90

4.3

7.3

3.4

2.3

0.82

0.69

1.9

0.81

0.82

0.09

4.3

7.6

4.2

?.7

0.92

0.86

2.4

1.2

1.0

0.25

4.7

8.2

4.3

2.7

1.0

0.90

2.7

1.2

1.1

0.44

4.8

8.5

4.4

2.9

1.0

0.95

2.5

1.2

1.2

0.32

5.2

9.5

5.4

3.5

1.2

1.1

3.2

1.7

1.2

0.80

5.7

10.8

6.8

4.4

1.4

1.4

3.8

2.2

1.2

0.70

7.2

waters, suggesting that Type A larvae might be referable
to S. (Sebastomus) exsul.

Distribution. — Eight CalCOFI cruises into the Gulf in
1956 and 1957 and an FAO cruise in March of 1972,
provide information on the distribution and abundance
of Type A larvae. Their latitudinal range, from Gonzaga
Bay south to Concepcion Bay, is more constricted than
that of S. cortezi, but they are not nearly so restricted to
the western side of the Gulf as are larvae of S. cortezi
(Fig. 16). Their heaviest concentration is around Tiburon
and Angel de la Guarda islands in the upper Gulf. Table
18 shows that spawning occurs during the cold-water
months with a possible peak in February.

Sebastes marinus (Linnaeus), Figure 17

Literature. — The numerous descriptions and illus-
trations of intraovarian and planktonic larvae of North
Atlantic Sebastes were cited above in the literature
review for the genus. Those of Taning (1961) are es-
pecially useful, since they incorporate observations on
chromatophores that can only be made in live material.
A series of illustrations from Taning's (1961) paper are
reproduced here. The larvae used for our study are from
the Dana collections and were identified by A. V. Tan-
ing, but Eschmeyer (pers. commun.) has suggested that
the series may contain some specimens of S. mentella
and that the identification should be considered ten-
tative until additional material is available. These were
measured to establish a table of morphometries, which to
our knowledge has not yet appeared in the literature. The
following description, which includes morphometric and
other previously overlooked characters, is intended as a
supplement to the extensive literature on S. marinus lar-
vae.

Distinguishing features.— At birth the larvae of S.

marinus are considerably larger than those of the eastern
Pacific species. A series of full-term intraovarian larvae
ranged in length from 6.7 to 7.2 mm. Also, they reach a
comparatively large size before undergoing a gradual
transformation into pelagic juveniles. In the measured
series the transition occurs at about 24 mm (Table 20).

26

Figure IS.— Larvae of SeboBtea Gulf of California Type A. A. 4.7 mm; B. 6.2 mm; C. 7.2 mm; D. 8.3 mm.

27

Figure 16.— Stations in the Gulf of
California at which larvae of
Seba»tes Gulf of California Type A
were taken on CalCOFI cruises
during 1956 and 1957.

Table 20. Measurerrents (run) of larvae of Sebastes marlmjs. (Spetlmens between dashed lines
are undergoing notochord flexion.)

Standard
length

Snout-
anus
distance

Head
length

Snout
length

Eye
diatneter

Body
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout. anal

fin

distance

7.1

2.2

1.2

0.33

0.51

1.3

0,18

0.26

...

7.3

2.9

1.8

0.50

0.66

1.4

0,50

0,55

...

...S.J._.

.... ?i?—

..!;§..

.?;5?-.

..P.-.'.3...

J_.4__

...o.-.S...

0.61 ..

8.S

3.2

2.1

0.55

0.78

1.6

0.68

0.66

0.04

4.5

9.0

3.8

2.5

0.73

0.87

1.9

0.75

0.74

0.06

4.1

9.3

3.9

2.5

0.68

0.86

1,8

0,74

0.75

0.09

5.3

9.6

4.0

2.7

0.75

0.91

1.9

0,82

0.80

0.12

5.6

10.0

4.5

2.8

0.93

1.0

2.2

1,0

0.82

1.15

5.7

10.6

4.8

3.2

1.0

1.1

2.3

1,1

0.87

0.18

6.0

11. 0

5.2

3.3

1.0

1.2

2,3

1,2

1.0

0.20

6.2

11.3

5.8

3.7

1.2

1.2

2,6

1,4

1,0

0.40

6.8

Jl.S

6.4

_4.0_

1.2

1.3

3.1

1_._5__

...v.".....

0.41 _

7.,4,._

"n'i"

..........

........

1,3

1.5

3.2

1.8

1,2

0-60

7.8

13.6

7.3

4.5

1.5

1.5

3.4

2.0

1,2

0.75

8.2

14.4

8.3

4.8

1.5

1.8

4.0

2.2

1,3

1.2

9,0

IS.O

8.8

4.9

1.6

1.7

3.9

2.3

1.3

1.2

9,4

16.6

9.8

5.8

2.0

1.9

4.3

2.6

1.3

1,5

10.3

17,2

10.0

5.8

1.7

2.2

4,2

3.2

1.4

1.8

10.5

18.4

10.2

5.8

1.7

2.3

4,6

3,3

1-4

1,9

10.6

19.3

11.0

6.6

2.1

2.4

4,9

3,5

16

2,1

11.7

20.5

11.7

6.7

2.0

2.4

4,9

3,8

1.5

2.3

12.3

22.3

12.8

7.1

2.1

2.6

5.2

4.2

1.7

2.8

13.4

•24.1

14.0

7.7

2.1

2.8

5.9

4.8

1.9

2.9

14.9

•26.0

15.4

8.7

2.4

3.0

6.5

5.8

2.1

3.6

16.2

•28.1

16.6

8.7

2.5

3.0

6,8

6.3

2.2

3.8

17.6

•30.4

18.2

9.4

2.9

3.2

7.7

6.8

2.3

4.3

19.3

•38.7

22.9

11.2

3.1

3.8

8.9

9.2

2,8

6.0

24.8

•Peljiglc juvenile.

The larvae are more slender than all but the slen-
derest of eastern Pacific Sebastes, such as S. jordani,
which they resemble in a number of other characters.

Body depth averaged 18% of body length in preflexion
larvae, 21% during flexion, and 25% in postflexion larvae
and early pelagic juveniles.

The gut is compact and relatively shorter than most
eastern Pacific Sebastes. In larvae less than 10 mm, the
relative snout-anus length is equal to or less than in
Sebastes jordani (Fig. 18). In developmental stages
larger than this, relative snout-anus length is inter-
mediate between S. jordani and the other eastern Pacific
species measured. Snout-anus distance increases from an
average of 35% of body length in preflexion larvae to 45%
during notochord flexion, and further to 58% in post-
flexion larvae and small pelagic juveniles.

The head is relatively small as in other species of
Sebastes. Head length averages 21% of the body length in
preflexion larvae, 29% in larvae undergoing flexion, and
32% in postflexion larvae and small pelagic juveniles.
Snout length averages 28% of head length in preflexion
larvae, increases to an average of 31% in later larval
stages, and then decreases slightly to an average of 29%
in small pelagic juveniles. Relative eye diameter is
greatest in preflexion larvae (mean of 40% of head
length) and then decreases to a mean of 35% of head
length in later developmental stages.

A prominent feature of S. marinus larvae is their
relatively short rounded pectoral fins. Fin length
averages 6% of body length before notochord flexion, 10%
during flexion, 17% in postflexion larvae, and 22% in
small juveniles. Relative fin length is less than in all
eastern Pacific species studied with the exception of S.
jordani which has similarly small pectorals (Fig. 19).
Pelagic juveniles of S. marinus have a slightly longer pec-
toral fin than in pelagic juveniles of S. jordani (Fig. 19).
The sequence of ossification of the pectorals and other
fins could not be determined because of calcium leaching
in the Formalin-preserved specimens.

The pigment pattern of S. marinus larvae is similar to
that of other North Atlantic Sebastes species and is also
similar to that of S. jordani of the eastern Pacific. New-
born larvae have a group of melanophores above the op-
tic lobes of the brain, a melanistic shield over the dor-
solateral surface of the gut, an embedded spot at the
nape, and a series of melanophores along the ventral and
dorsal midlines of the tail. The ventral row begins at the
4th to 7th postanal myomere and extends to the 19th to
22nd postanal myomere. In a sample of 18 full-term in-
traovarian larvae, the total number of melanophores in
the ventral row ranged from 11 to 24 with a mean of

18.0 ± 4.07 SD. The dorsal row is shorter, beginning on
the 10th to 15th postanal myomere and ending on the
18th to 22nd postanal myomere; the number of
melanophores ranged from 8 to 21, with a mean of

13.1 ± 3.16 for the 18 specimens.

The ventral midline series of S. marinus is shorter and
contains fewer melanophores than in larvae of the other
North Atlantic species and in S. jordani of the Pacific. In
120 newborn specimens of the American form, the num-
ber of ventral melanophores ranged from 26 to 42 and in
three newborn specimens of S. viviparus from the Faeroe
Islands the number ranged from 18 to 29. In 15 specimens

28

Figure 17. — Developmental series of Sebastes marimu from Taning (1961). A. 6.8-mm larva; B. lO.S-mm larva; C. 15.7-min larva;

D. 20.9-nun larva; E. 27.0-inni pelagic juvenile.

29

■ib

r-

«

20

-

o 4> Sebostes

mocdonaldi

*

^

*

*

• • S

" * s.

fosciolus

monnus

A

15

-

• ♦ s.

D « S

■ « s

oaucispinus

levis

lordani

o ♦ * ■••

* <. .■

10
5

n

>

o

o **

1 . 1

1

1

20 24 26

STANDARD LENGTH (mm)

Figure 18. — Relation of snout-anus length to body length in developmental stages of

Sebostes spp.

o

* Sebos

tes mocdonoldi

•

• S

fosciotus

^

* s

monnus

A

♦ s

poucispinus

O

« S

levis

E

E

■

• s

lordani

<

O

.*■

» .d*^ •erf'

20 21 26

STANDARD LENGTH (mm)

Figure 19. — Relation of pectoral fin length to body length in developmental stages of

Sebaatet spp.

of S. jordani, the number of ventral melanophores ranged
from 21 to 27 with a mean of 23.7 ± 1.75 SD. Larvae of S.
marinus differ also from those of the American form and
S. viviparus in lacking the one to several caudal
melanophores present in these latter species.

The dorsal and ventral midline pigment series of S.
marinus larvae have opposite fates. The ventral series
becomes embedded in the developing musculature and
obscured by the developing anal fin. At the completion of
notochord flexion only those melanophores posterior to

the anal fin are visible. The dorsal series is augmented
anteriad and in 9-mm larvae divides into two streaks,
one on each side of the developing dorsal fin. When the
larvae reach 14.5 mm the streaks extend anteriad along
the entire dorsum.

In addition to the initial pigment pattern,
melanophores are added gradually to the head region. In
9-mm larvae a pair of melanophores appears on the ven-
trolateral surface of the medulla, the olfactory region of
the brain becomes covered with a patch of melanophores,

30

and a pair of melanistic streaks appears along the
premaxillaries. At about 11.0 mm, a melanistic patch ap-
pears on the opercle and enlarges with further develop-
ment. At about 12.5 mm, a blotch appears at the sym-
physis of the lower jaw. In 15-mm larvae, blotches appear
around the nares, ventroposterior to the eyes, and at the
isthmus. At about 17.0 mm, melanophores cover most of
the dorsal and lateral surfaces of the head.

On the body the initial pigment pattern is augmented
by the appearance of melanophores at the posterior edge
of the hypural plates in 9-mm larvae. At the completion
of notochord flexion, the posterior edge of each hypural
plate is outlined by a pigment streak. At about 10.0 mm,
deeply embedded pigment begins to form along the dor-
sal aspect of the more posterior vertebral centra. This
spreads anteriorly and at about 16.0 mm the dorsal as-
pect of the entire vertebral column is covered. At about
13.5 mm a line of melanophores forms along the pos-
terior segment of the lateral line and extends progres-
sively anteriad with further larval development. Also,
each of the distal radial elements of the dorsal fin
develops a melanistic spot as do the distal radials of the
anal fin in 15-mm larvae.

The epaxial region of the trunk begins to be covered
with melanophores in 16-mm larvae. These are concen-
trated along the myosepta. At about 24.0 mm the entire
half of the trunk above the lateral line is covered, mark-
ing the transition to the pelagic juvenile phase. The pec-
toral and pelvic fins develop no melanistic pigmentation
in larvae or pelagic juveniles.

Distribution. — The geographic and bathymetric dis-
tribution of Sebastes in the North Atlantic is the subject
of an enormous literature and need not be discussed here.
Much of the information on distribution of adults is sum-
marized in Templeman (1959) and in the proceedings of
an international symposium on redfish (Templeman
1961), although most of his references to S. marinus men-
tella refer to S. fasciatus (Eschmeyer, pers. commun.).
The symposium proceedings also contain a summary of
information on distribution of larvae (Einarsson 1961).

Sebastes viviparus (Kr»(yer), Figure 20

Literature. — Literature references on S. viviparus lar-
vae were cited in the literature review for the genus. In
the earlier literature, larvae of S. viviparus were
sometimes confused with S. marinus. Templeman and
Sandeman (1959) have reviewed this literature and at-
tempted to correct the errors. The two most useful papers
on S. viviparus larvae are by Taning (1961) and
Einarsson (1960). In these papers, larvae of S. viviparus
and S. marinus are described and compared with em-
phasis on the differentiating characters of pigmentation
and spination.

A series of larvae up to 10.6 mm was obtained from the
collections of the Dana Expeditions and measured to
provide a table of morphometries for comparison with
other North Atlantic species (Table 21). This is

Table 2\ . MeasurerMnts ("*") o^ larvae of Sebastes vlylparus. (Specimens between dashed
lines are undergoing notochord flexion.)

Standard
length

Snout-
anus
distance

Head
length

Snout
length

Eye
diameter

Body
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin
distance

5.4

2.0

1.2

0.28

0.55

1.0

0.35

0.43

5.9

2.Z

1.2

0.37

0.57

1.2

0.53

0.44

—

...

6.5

2.4

1.3

0.32

0.59

1.2

0.40

0.41

6.8

2.3

1.3

0.31

0.60

1.2

0.45

0.46

7.0

2.8

1.8

0.48

0.72

1.5

0.57

0.56

....'.■.3..J

...2.5...

...h}..

..il.M..

..SM...

.i.i...

..S.ii....

..O.il...

...-.-.-—

.--.-...,

;.e

3.0

1.9

0.50

0.80

1.7

0.62

0.69

-„

8.0

3.2

2.0

0.58

0.82

1.7

0.62

0.68

3.7

8.5

3.7

2.3

0.69

0.92

1.8

0.78

0.80

0.13

4.6

9.0

3.9

2.6

0.78

1.0

2.1

0.90

0.95

0.19

5,1

9.4

4.2

2.8

0.86

1.0

2.2

0.92

1.0

0.17

5.3

9.9

4.5

2.9

0.97

1.1

2.3

1.1

1.0

0.18

5.7

10.4

4.8

3.2

1.0

1.2

2.4

1.2

1.0

0.28

6.1

__jp^6___

5.2

...hi..

._J^p_._

..h?....

.2.6...

..hi.....

-PJP...

..P.M..

i.Z....

presented with a brief description that incorporates
previously unreported characters together with those of
Taning (1961)

Distinguishing features. — Sebastes viviparus larvae
are born at a size 1 to 2 mm smaller than S. marinus lar-
vae. Taning (1961) illustrated a 5.8-mm full-term in-
traovarian larva of S. viviparus; the smallest planktonic
larvae in our series was 5.4 mm. Other developmental
events also occur at a smaller size in S. viviparus com-
pared with S. marinus. Notochord flexion begins at
about 7.8 mm in our series and is completed at about 10.6
mm. In S. marinus it begins at about 8.5 mm and is com-
pleted at about 11.8 mm. Einarsson (1960) and Taning
(1961) pointed out that development of the head spines,
particularly the preopercular series, is at a relatively ad-
vanced state in S. viviparus larvae of the same size as
those of S. marinus. The size at transformation into the
pelagic juvenile stage cannot be ascertained from our
series since the largest larva is 10.6 mm. There are no
literature references to size at transformation for this
species.

Developmental changes in body proportions follow
those of S. marinus with some apparent differences in
mean values for the principal larval stages. Means for
relative eye diameter, body depth, and pectoral fin base
depth are greater than in S. marinus for preflexion lar-
vae and larvae undergoing notochord flexion (Table 22).

The pattern of melanistic pigmentation is similar to
that of S. marinus; however, the presence of one or more
caudal melanophores in S. viviparus larvae separates
this species from S. marinus (Templeman and
Sandeman 1959; Einarsson 1960; Taning 1961). The in-
completeness of our series precludes a thorough com-
parison of pigment pattern in S. viviparus with those of
other North Atlantic species, but, we have discovered
some previously unreported differences.

Pigment features develop in smaller sized larvae of S.
viviparus compared with S. marinus. Several heretofore
unreported features begin to appear in larvae about 6.0
mm long. A group of melanophores form at the tip of the
snout and remain prominent up to the termination of our
series at 10.6 mm. Although a melanophore may be pres-
ent in this region in occasional specimens of S. marinus

31

B

F1ffure20.— Larvaeof5eba«te8 vivipartM from Taning (1961). A. 5.8 mm; B. 8.9 mm; C. 10. 1 mm;D. 13.5 mm.

TABLE 22. Morphometric comparison of larvae of three species of Sebastes from the North Atlantic. Mean percentage and range of body proportions are given
for preflexion (A), flexion (6). and postflexion (C) larvae. Measurements in rrm.

Sebastes

Snout-
anus distance

Head lenqth

Snout length

Eye diameter

Body depth
SL

Pectoral
fin length

Pectoral fin

base depth

SI

Pelvic fin
length

Snout-anal
fin distance

SL

SL

Head le

nqth

Head

5L

SI

SL

7+50

Ranqe

J + SD

Ranqe

5 ± SO

Ranqe

5 + SD

Ranqe

X + SD

Range

; 1 SD

Range

; 1 SD

Range

; 1 SD

Range

X 1 SD

Ranqe

37.2+2.6

34-40

22.0+2.6

19-26

27.6+2.9

23-31

42.8+3.3

38-46

18.413.5

13-22

6.210.4

6-7

6.711.5

6-9

- -

-

- -

-

fasciatus

46.0±?.0

44-48

29.0t2.0

27-31

30.0t2.6

28-33

37.7+3.1

35-41

24.711.1

24-26

8.012.0

6-10

8.3+0.6

8-9

1.610.6

1-2

57.313.1

54-60

55.2+3.2

51-59

32.7±1.2

31-34

31.0+2.1

27-33

34.811.6

33-37

28.2*1.7

26-31

15.312.0

13-18

8.710.5

8-9

6.813.4

3-12

60.213.1

58-61

35.7+4.5

31-40

21.3t4.0

17-25

27.7+0.6

27-28

40.3+3.1

37-43

18.011.0

17-19

5.712.3

3-7

6.712.3

4-8

- -

-

marinus

45.1+5.0

38-54

29.2±2.9

25-34

30.3+3.1

26-36

34.6tl.6

32-37

21.412.2

19-26

9.911.8

8-13

8.210.4

8-9

1.711.1

0.5-4

56.314.7

53-63

56.9+1.7

54-59

33.2t0.9

32-35

31.2+1.7

29-34

36.212.2

33-40

25.1+1.4

23-28

16.8+2.0

14-19

8.3+0.7

7-9

9.112.4

5-13

61.011.5

58-63

37.5*2.6

34-40

22.2+3.1

19-26

26.0+2.8

24-31

44.0+3.6

39-48

19.5+1.4

18-21

7.311.2

6-9

7.511.0

6-9

- -

-

- -

-

viviparus

43.7+3.4

38-49

28.2+2.7

24-31

30.0+2.0

26-33

38.6+2.2

36-42

22.611.3

21-25

9.911.5

8-12

9.611.1

8-11

2.210.4

2-3

55.414.5

46-59

32

and the American form, larvae of these species lack the
obvious patch of melanophores that is present in S.
viviparu.s. Gut pigmentation is slightly different in S.
viviparus larvae compared with the other species. The
dorsolateral surfaces of the gut are covered with a
melanistic shield as in the other species; however, in S.
viviparus a covering of less densely distributed
melanophores extends ventrad over the ventral surface of
the gut. The ventral surface of the gut is unpigmented at
comparative larval stages in S. marinus and S. fas-
ciatus.

The most outstanding difference in pigmentation is in
the pectoral fins. Sebastes marinus and S. fasciatus lack
pectoral fin pigment throughout the larval period while
S. viviparus larvae have distinctive pectoral fin pig-
ment. At about 6.0 mm, the medial surface of each fin
base begins to develop a covering of melanophores, which
remains throughout the larval stages available to us.
Also, a series of fine melanophores forms along the bases
of the rays and may extend distally for a short distance
between the rays in some specimens. This character
should be of considerable help in separating larvae of S.
viviparus from those of other North Atlantic species.

The few early larvae of S. viviparus available to us
precludes analysis of the number of melanophores in the
ventral midline series. Of the three smaller larvae
counted, the ventral row began on the 3rd or 4th postanal
myomere and terminated on the 19th or 21st myomere.
The number of melanophores ranged from 18 to 29 with a
mean of 25. This suggests that the mean number of
melanophores is greater in S. viviparus than in S.
marinus, a supposition that can only be verified by anal-
ysis of an adequate sample of intraovarian larvae.

Distribution, — See Templeman (1961).

Sebastes fasciatus (Storer), Figure 21

Literature. — Bigelow and Welsh (1925) briefly de-
scribed and illustrated a series of Sebastes larvae from
the Gulf of Maine. Barsukov (1968, 1972) and Barsukov
and Zakharov (1972) have established that the common
form of this region is specifically distinct and should be
referred to as S. fasciatus Storer. This is confirmed by
Eschmeyer's recent work on gas bladder myology (cited
in Hallacher 1974). A larval series oi Sebastes (Table 23)
collected off New England from lat. 38°52'N to lat.
44°22'N by the NMFS Laboratory, Narragansett, is de-
scribed briefly below and compared with those of S.
marinus and S. viviparus.

Distinguishing features. ~\t would appear that lar-
vae of this form are born at a smaller size than those of S.
marinus (Tables 17, 20); however, judging from the ap-
pearance of our small specimens of S. fasciatus, this dif-
ference could be a result of shrinkage in preservative.
Larvae of both forms begin notochord flexion at about 8.5
mm; however, flexion is completed at 10.0 mm length in
S. fasciatus and at about 11.8 mm in S. marinus. Infor-

mation on the size of transformation is not available
since the largest larva is 17.1 mm.

Developmental changes in body proportions follow
closely those of S. marinus and need not be repeated here
(Tables 20, 23). One feature which appears to differ is
relative body depth. Table 22 shows that mean relative
body depth is greater during the three larval phases in S.
fasciatus than in S. marinus. Mean relative eye diameter
is greater in S. fasciatus than in S. marinus for preflexion
larvae and larvae undergoing flexion, but is less in post-
flexion larvae. Mean relative pectoral fin length is
greater in S. fasciatus than in S. marinus in preflexion
larvae but is less during and after flexion.

Pigmentation is similar to that of S. marinus; however,
analysis of our specimens has revealed some differences
which have not been previously reported. As in S.
marinus larvae, the initial pigment pattern consists of a
group of melanophores above the brain, a gut shield, an
embedded spot at the nape, and a series of melanophores
along the dorsal and ventral midlines. There appears to
be a difference in the number of melanophores com-
posing the ventral midline series of S. marinus and S.
fasciatus larvae. Ventral melanophores were counted in
10 late-stage intraovarian larvae from each of 12 females
of S. fasciatus taken in the Gulf of Maine. The mean
numbers of melanophores for the 12 samples were as fol-
lows: 31.4, 32.2, 32.6, 32.8, 32.8, 33.3, 34.9, 35.2, 35.5,
37.5, 38.3, 40.4. The range for all specimens was 26 to
42. In a sample of 18 S. marinus larvae of comparative
developmental stage from Greenland, the mean was 18.0
with a range of 11 to 24. Also, the ventral pigment line is
relatively longer in S. fasciatus, beginning on the 1st to
4th (mean of 2.9 ± 0.40 SD for 112 specimens) postanal
myomere and ending on the 19th to 23rd (mean of
21.3 ± 0.85) postanal myomere. In the sample of S.
marinus the series begins on the 4th to 7th (mean
5.4 ± 0.78 for 18 specimens) postanal myomere and ends
on the 19th to 22nd (mean of 20.9 ± 0.86) postanal
myomere. The dorsal midline row appears to contain
fewer melanophores in S. fasciatus compared with S.
marinus. Mean counts for the 12 samples are: 3.0, 5.4,
6.6, 6.9, 7.2, 7.2, 7.3, 7.6, 7.9, 8.0, 8.6, 9.1. In the sample
of S. marinus the mean was 13.1 with a range of 8 to 21.
In the samples of S. fasciatus the dorsal series began on

Table 23. Measurements (mn} of larvae of Sebastes fasciatus. {Spectmens between dashed
lines are undergoing notochord flexion.)

Standard
length

Snout-
anus
distance

Head

length

Snout

length

Eye

diameter

8ody
depth

Pectoral
fin
length

Pectoral

fin base
depth

Pelvic

fin

length

Snout-anal

fin
distance

5.8

2.0

1.1

0.25

0.51

1.0

0.3S

6.2

2.4

1.3

0.36

0.58

l.I

0,36

0.40

7.1

2.7

1.6

0.44

0.71

1.4

0.40

0.42

7.5

2.6

1.6

0.44

0.66

1.5

0.42

0.43

8.Z

3.3

2.1

0.65

0.80

1.8

0.61

..9.??...

..P--P7..

-...4j2...

8.5

3.7

2.3

0.77

0.95

2.0

0.5S

0.67

0.11

4.6

9.2

4.2

2.7

0.75

1.0

2.2

0.76

0.80

0.15

5.5

10.0

4.8

1.1

_J.1__

""I'r"'

2.6

"Co"

1.0

1-5

'""i"o""

0.25
0.42

____5.8 __

6.3

10.8

5.7

11.?

5.7

3.6

1.1

1.2

2.9

1-4

t.o

0.37

6.8

12.0

6.4

3.8

1,2

1.4

3.4

1.7

1.1

0.55

7.1

13,3

7,6

4.4

l.Z

1.6

3.9

2,2

1.2

1.1

8.1

14,4

8.5

4.9

1.6

1.7

3,9

2.3

1.2

1.3

8.8

17,1

9.9

5.8

1.8

1.9

4.8

3.1

1.4

2.1

10.5

33

. --1

— ^

•> V «

iSS^vS^if:';'' ■■

;-^W

-^^^^

Figure 21.— Larvae of 5eb<Mte8 fasciatus (A-D) from Bigelow and Welsh (1925) and a larva of Sebastes capensis (E). A. 6 mm; B.

9 mm; C. 12 mm; D. 20 mm; E. 7.2 mm.

34

the 7th to 14th (mean of 10.9 ± 1.35 for 111 specimens)
postanal myomere and ended on the 14th to 22nd (mean
of 19.1 ± 1.19 for 110 specimens) postanal myomere. In
S. marinus the series began on the 10th to 15th (mean of
12.7 ± 1.41) postanal myomere and ended on the 18th to
22nd (mean of 20.3 ± 1.08) postanal myomere. In con-
trast to the ventral row in which the number of con-
stituent melanophores is fixed during late intraovarian
life, the dorsal row is gradually augmented before and
after birth. The fact that the sample of S. marinus lar-
vae was further advanced than any of the broods of S.
fasciatus studied may account for the greater number of
dorsal melanophores in the former. Consequently, a de-
tailed study of comparable pre- and postnatal larvae of
the two species would be necessary to evaluate the utility
of this character in separating their larvae.

A well-known difference in the initial pigment pattern
of the two larval types is the presence of one or more
melanophores at the caudal region in S. fasciatus and the
absence of caudal melanophores in most larvae of S.
marinus (Templeman and Sandeman 1959). Caudal
melanophores were present in all intraovarian larvae of
S. fasciatus studied. The frequency was as follows: 40 lar-
vae had 1 melanophore, 64 had 2, 14 had 3, and 2 had
4.

The subsequent development of pigment pattern in
larvae of S. fasciatus is similar to that in S. marinus, al-
though a number of differences are apparent. The ven-
tral midline series becomes embedded gradually but
some melanophores remain along each side of the base of
the anal fin even in our largest larvae. In S. marinus lar-
vae the developing anal fin obscures the ventral midline
series and the remnants are visible only posterior to the
fin in larvae longer than 11.8 mm. The dorsal midline
series extends anteriad and splits into two lines with the
developing dorsal fin between. The entire dorsum is
spanned when the larvae are about 10.8 mm. In S.
marinus larvae the entire dorsal midline is not spanned
until the larvae reach about 15.0 mm.

The appearance of pigment characters at a smaller size
(usually 1 to 3 mm smaller) in larvae of S. fasciatus com-
pared with S. marinus larvae is a general developmental
feature. For instance, pigment appears above the olfac-
tory region of the brain in S. fasciatus larvae about 7.0
mm long and in S. marinus larvae about 9.3 mm
long.

Another difference is the absence of certain pigment
characters in S. fasciatus larvae and their presence in S.
marinus of comparative size. Pigment does not appear on
the isthmus, ventroposterior to the eye, and on the dis-
tal radials of the anal fin in S. fasciatus larvae of the size
range studied, while in S. marinus larvae, these pigment
features appear at about 15.7 mm. A more complete
series of S. fasciatus larvae will be necessary to ascertain
at what size these features appear or whether they ap-
pear at all.

Distribution. — Information on the geographic and
bathymetric distributions of S. fasciatus is given by
Templeman (1959).

Sebaates capensia (?) (Gmelin), Figure 21E

Literature. — Larvae of Sebastes from the Southern
Hemisphere have not been previously described. The
taxonomic status of this genus in the Southern Hemi-
sphere is confusing, and Chen (1971) in his review of the
subgenus Sebastomus, to which the southern forms be-
long, lumped the 11 nominal species into one. Eschmeyer
and Hureau (1971) and Eschmeyer (pers. commun.),
after examining specimens in South American museums,
feel that there are at least three species off Chile. A total
of 13 larval specimens from off Chile are available — five
from the Marchile VI Expedition, six from Piquero TV,
and two large specimens dip netted under a night-light in
Valparaiso Harbor by Richard McGinnis of Pacific
Lutheran University. Although some of these specimens
are in poor condition and the series is incomplete, a brief
description of them accompanied by an illustration
serves as a basis for comparison with Sebastes from other
regions of the world's oceans.

Distinguishing features, — Newborn larvae are small
(about 3.8 mm), a feature which agrees with Sebastes
larvae of the northeastern Pacific, and particularly the
subgenus Sebastomus (Fig. 1). The larvae attain a large
size, since the 19.6-mm dip net specimen is just begin-
ning to transform.

Morphometric information on this species is incom-
plete because of the brevity of the developmental series
(Table 24). The proportions generally follow those of
other species, however, the depth of the pectoral fin base
appears to be slightly greater in this species than in
others measured, with a mean of 12.4'^ of the body
length and a range of 8 to 17% for the seven larvae
measured.

Pigmentation is similar to many of the Sebastomus
species of the northeastern Pacific. Initially, the larvae
have a spot above the nape, a spot on each side of the
medulla, a group of melanophores at the symphysis of
the lower jaw, a melanistic shield dorsolaterally on the
gut, a group of melanophores on the ventral surface of the
gut, and a series of melanophores along the ventral mid-
line of the tail. The number of melanophores in the ven-
tral midline ranged from 15 to 19 in four small larvae.
The pectoral fins are distinctly pigmented with a group
of melanophores on the medial surface of the fin base and
melanophores distributed over the blade of the fin.

One or more melanophores lie above the optic lobes of
the brain in larvae 4.6 mm and longer. As the pectoral fin

Measurements (nm) of larvae of Sebastes capens Is. (Specimens between dasbed lines
are undergoing notocboro Tlexion.)

Standard
length

Snout-
anus

distance

Head
length

Snout
length

Eye
diameter

Body
depth

Pectoral
fin
length

Pectoral
f1n base
depth

Pelvic

fin

length

Snout -anal

fin

distance

3.8

1.5

0.80

0.19

0.35

0.85

0.33

0.32

...

...

4.6

t.fi

1.1

0.36

0.43

1.2

5.5

2.6

1.8

0.60

0.59

1.7

0.62

0.72

0.05

...

6.2

2.6

1.9

1.8

0.77

0.75

0.06

3.7

7.0

3.7

2.3

0.83

0.93

2.6

1.3

1.1

0.28

4.3

7.2

4.0

2.7

0.93

0,94

2.7

1.3

1.2

0.65

4.3

14.8

6.9

5.6

1.5

1.7

5.5

3.8

1.7

2.5

9.4

19.6

12.0

7.1

2.1

2.1

6.,

5.3

2.0

3.7

12.8

35

rays differentiate, the melanophores become restricted to
the interradial membrane and the distal margin of the
pectoral fin develops a melanistic outline. The 7.2-mm
larva has added a melanophore above the olfactory lobes,
a spot on the ventrum at the juncture of the cleithra, and
a covering of melanophores on the pelvic fins.

The gap in the series between 7.2 and 14.8 mm
prevents study of pigment changes in this size range. The
14.8- and 19.6-mm specimens are pigmentless except for
a wide band of melanophores on the caudal peduncle. In-
terestingly, many and possibly all species oi Sebastomus
from the northeast Pacific have a caudal peduncle band
at this stage.

Fin counts on the two dip net specimens (dorsal, XIII,
13; anal, III, 6; pectoral, 18) fit known meristics for S.
capensis. A count of 42 lateral line pores on the 19.6-mm
specimen falls within the range for S. capensis.

Distribution. — Chen (1971) summarized the dis-
tributional information on this species.

Se6a«te8— Northwestern Pacific Species, Figure 22

Literature. — According to Chen (pers. commun.)
there are 31 species of Sebastes in the northwestern Pa-
cific. Life history series have been published on eight of
these.

Fujita (1957) reared larvae of S. pachycephalus nig-
ricans which had been extruded spontaneously by a cap-
tive pregnant female. The larvae were fed brine shrimp
nauplii and kept at 16.4° to 18.8°C for 1 mo. The larval
series was described and illustrated.

Fujita (1958) obtained a series of developing embryos
of S. oblongus from two pregnant females maintained in
an aquarium at 13.5° to 18.0°C. Larvae from one female
were reared at 16° to 18°C on a diet of brine shrimp
nauplii for 30 days until fin formation was completed.
The series of developing embryos and larvae was de-
scribed and illustrated.

Uchida et al. (1958) described and illustrated three
specimens of S. hubbsi taken from plankton hauls.

Shiokawa and Tsukahara (1961) obtained a series of
embryos of S. pachycephalus pachycephalus from cap-
tive females and then reared the larvae for 25 days at
15°C on a diet of brine shrimp nauplii. The embryos and
larvae were described and illustrated as were demersal
juveniles up to about 25-mm length netted from shallow
water.

Harada (1962) described and illustrated a series of S.
inermis that included late preextrusion larvae taken
from pregnant females, larvae collected by plankton net,
and juveniles up to about 60 mm in length.

Takai and Fukunaga (1971) followed development of
embryos in a captive female of S. longispinus and reared
the larvae for 30 days on a diet of brine shrimp and Tig-
riopus nauplii. The embryos and larvae were described
and illustrated.

Sasaki (1974) removed late preextrusion larvae from
freshly killed females of S. schlegeli, S. steindachneri,
and S. taczanowskii. He summarized morphological and

pigmentary characters of 20 specimens of each species
and illustrated a specimen of each.

Distinguishing features. — Larvae of S. oblongus
hatch at a total length of 7.25 mm or 6.5 mm notochord-
al length.' This is 1 mm larger than any eastern Pacific
species described and slightly less than the range of S.
marinus of the Atlantic. The most outstanding feature of
S. oblongus larvae is the pattern of melanistic pigment,
which begins to form well before hatching. Melanophores
first appear as a shield over the dorsal surface of the gut,
then patches form on the dorsal surface of the head, on
the trunk above the gut, and as a band on the tail. At
hatching, melanophores are continuous from the head to
the broad tail band and they also appear on the ventral
surface of the gut. The pectoral fins are unpigmented.
With further development, pigment forms on the
preopercular region of the head, along the jaws, the bases
of the pectoral fins, and the tail band extends anteriorly
and posteriorly. At notochord flexion the larvae are
covered with melanophores except for the distal half of
the pectoral fin blade and the caudal region (Fig. 22A).
Larvae of S. oblongus differ from eastern Pacific and At-
lantic species in their extraordinarily heavy pig-
mentation and in not developing the series of ventral
midline melanophores on the tail.

Larvae of S. longispinis are spawned at about 5.3 to 5.5
mm notochordal length which is equivalent to S. jor-
dani. the largest eastern Pacific species at hatching. In-
terestingly, the brood of S. longispinis studied by Takai
and Fukunaga (1971) hatched as embryos within the
ovary at about 3.1 mm notochordal length. Although this
may have resulted from stress during captivity, it fur-
ther strengthens the view that hatching occurs within the
ovary in Sebastes and not after extrusion as stated by
some authors (e.g., Morris 1956; Waldron 1968). The
development of the pattern of melanistic pigmentation is
almost identical to that in S. oblongus (Fig. 22B). The
only difference appears to be in the absence of pigment
on the distal portion of the pectoral fin base and on the
blade of the fin.

Larvae of S. hubbsi represent a third member of this
heavily pigmented group of Japanese Sebastes. The
smallest of the three planktonic specimens described by
Uchida et al. (1958) was a newborn larva 4.2 mm in
notochordal length, indicating that larvae of this species
are born at a length 1 mm smaller than larvae of S.
longispinis and about 3 mm smaller than S. oblongus.
The pigment pattern develops almost identically to that
in the other two species, but differs chiefly in the sparse
pigmentation of the pectoral fin base and in the develop-
ment of evenly distributed minute melanophores on the
blade of the pectoral fin (Fig. 22C).

'It is customary in Japanese descriptions of fish larvae to give body
length as total length, including the finfold. Since we use notochordal
length in preflexion larvae, the total lengths were converted to notochord-
al length by measuring notochordal and total lengths on the illustrations
and multiplying the resultant conversion factor by the actual total length
given for the specimen.

36

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37

A second type of Sebastes larva is represented by the
subspecies S. p. pachycephalus and S. p. nigricans. Lar-
vae of the two subspecies are extruded at lengths of 5.5 to
6.4 mm and 6.2 to 6.3 mm notochordal length, respec-
tively, slightly larger than in eastern Pacific species.
They also develop a striking pattern of melanophores.
The first pigment to appear in embryos is a patch over
the dorsal surface of the gut and a patch directly above
this over the trunk. With further embryonic develop-
ment, a large patch forms on the dorsal surface of the
head and the base and blade of the pectoral fin is covered
solidly with melanophores. As in the S. oblongus group,
larvae of S. pachycephalus do not develop the series of
ventral midline melanophores that is present in eastern
Pacific and Atlantic Sebastes. At hatching, the heavy
trunk band is augmented and melanophores have spread
to the lateral and ventral surfaces of the gut. With fur-
ther development, melanophores are added to the oper-
cular region and the pectorals become solidly black.

The two subspecies differ in the pigmentation of the
trunk and tail. Larvae of S. p. pachycephalus augment
the anterior trunk band throughout larval development
but develop no pigmentation on the posterior trunk and
tail (Fig. 22D). Late in the larval period, two wide bands
develop on the tail, one below the soft dorsal fin and one
at the caudal peduncle. In preflexion larvae of S. p. nig-
ricans, a line of melanophores extends posteriorly from
the heavy trunk band to the caudal region (Fig. 221). In
later larval stages, melanophores are added to the lateral
trunk and tail to form a continuous mass of pigment
along the lateral body surface. The pelvic fins are solidly
pigmented in both subspecies.

A third type of pigment pattern in Japanese Sebastes
larvae is similar to that found in some species of the
eastern Pacific and Atlantic. Late intraovarian larvae of
S. steindachneri (Fig. 22E), S. taczanowskii (Fig. 22F),
and S. schlegeli (Fig. 22G) all have a row of
melanophores on the ventral midline of the tail, an op-
posing row along the dorsal midline, a patch on the dor-
sal surface of the brain, and a solid shield over the gut.
They differ in the number of melanophores in each of the
rows. Mean values for the number in the vental row of S.
steindachneri, S. taczanowskii, and S. schlegeli are 16.3,
9.9, and 9.5, respectively, and for the dorsal row 18.7, 9.3,
and 10.3, respectively. Size ranges in notochordal length
for full-term larvae of the three species are 3.4 to 4.6 mm,
4.2 to 5.2 mm, and 4.8 to 6.8 mm, respectively.

A fourth species that falls into this group is S. iner-
mis. Larvae are born at about 4 to 5 mm in length and
Harada (1962) shows only dorsal head and gut pigment
at this stage. His illustrations of 6- to 10-mm larvae show
a short row of ventral and dorsal midline melanophores,
but no numbers are given (Fig. 22H). No pectoral fin pig-
ment is present on this species nor on the preceding
three.

Helicolenua Goode and Bean

Literature. — Larvae of Helicolenus dactylopterus
have been described and illustrated by a number of

workers. Fage (1918) described a series of larvae 5.5 to 14
mm long from the Mediterranean Sea and illustrated the
largest and smallest individuals of the series. Sparta
(1942, 1956) described more completely the larval and
early juvenile stages of H. dactylopterus from the
Mediterranean and illustrated numerous larval stages
and some juveniles. Haigh (1972) described the os-
teological development of a larval series from off South
Africa. A series of excellent illustrations of H. dactylop-
terus are included with Taning's (1961) description of
redfish larvae from the North Atlantic.

Distinguishing features.— Larvae of Helicolenus can
be differentiated from those of other scorpaenid genera
by a combination of characters. Like Sebastes, larvae of
H. dactylopterus have a small pectoral fin base (Table
3). Depth of the fin base averages 11% of the body length
in larvae before notochord flexion, 12.5% during flexion,
and 12% after flexion is completed. The range for the en-
tire larval period is 9 to 14%. The blade of the fin is short
and rounded. Fin length averages 11% of the body length
in preflexion larvae, 15% during flexion, and 18% fol-
lowing flexion. Another distinctive feature is the mass of
spongy tissue which forms at the dorsal midline of the
trunk. It appears within the dorsal finfold in 4-mm
larvae in the region to be occupied by the spinous
dorsal fin. In later stages the spinous dorsal rays form
within the mass and erupt from it. Such tissue has
not been described for larvae of any other scorpaenid
genus.

Larvae of the species of Helicolenus from the eastern
Pacific off Chile, H. lengerichi, were not available for
study. For comparative purposes, a brief description of a
series of the North Atlantic species, H. dactylopterus, is
given below and accompanied by the illustrations of Poul
Winther (Taning 1961).

Helicolenus dactylopterus (Delaroche), Figure 23
Literature. — See above.

Distinguishing features. — Planktonic eggs of H. dac-
tylopterus have not been described, and there is uncer-
tainty about the mode of reproduction of this species.
Krefft (1961) reported that ovaries of reproductively
mature females contained developing embryos em-
bedded in a gelatinous matrix. Thus, Helicolenus may be
ovoviviparous as in the subfamily Sebastinae or may pro-
duce a gelatinous egg balloon as in many genera of the
Scorpaeninae. Krefft (pers. commun.) has suggested
that the ovoviviparity may be facultative. Larvae are
small at hatching as in the Scorpaeninae; Sparta (1942)
illustrated a 2.8-mm larva that has already utilized its
yolk and is morphologically similar to Scorpaena larvae
of that size.

Body depth is moderate and increases throughout the
larval period, from an average of 29% of the body length
in preflexion larvae, to 33% in larvae undergoing flexion,
to 37% in postflexion larvae (Table 25). The gut is com-
pact. Snout-anus distance averages 49% of the body

38

B

Figure 23.— Larvae of Helicolenus dactylopterus from Taning (1961). A. 3.6 mm; B. 5.6 mm; C. 10.0 mm; D. 19.0 mm.

39

standard
length

Snout-
disunce

Head
length

Snout
length

Ejre
diaineter

Body
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin

distance

3.2

1.5

0.95

0.28

0.29

0.81

0.34

0.35

...

3.8

l.S

0.45

0.35

1.2

0.40

0.35

4.6

2.3

0.53

0.43

1.3

0.52

0.55

5.0

2.6

0.63

0.47

1.4

0.60

0.60

...i^...

.. 2.8

0.62

0.58

1.8

0.71

0.65

0.04

3.5

6.0

3.0

0.82

0.63

2.0

0.80

0.69

0.08 1

3.6

6.2

3.0

0.82

0.62

1.9

0.90

0.74

0.10

3.6

6.6

3.1

0.80

0.65

2.2

0.92

0.79

0.14

3.8

6.S

3.4

0.96

0.74

2.3

1.1

0.90

0.15

4.0

6.9

3.3

0.92

0.77

2.3

1.1

0.92

0.15

4.0

7.9

4.3

1.1

0.78

2.7

1.2

1.0

0.32

5.2

S.1

4.7

1.2

1.0

2.6

1.3

1.1

0.50

5.0

S.6

4.8

1.2

1.0

3.2

1.5

1.2

0.70

5.4

9.8

5.8

1.2

1.2

3.6

1.7

1.2

1.1

6.2

10.7

6.2

1.3

1.3

4.1

2.1

1.3

1.2

6.7

15.4

9.1

6.2

2.2

1.7

5.6

3-1

1.6

2.1

9.6

length in larvae up to notochord flexion and thereafter
increases to an average of 58%. Head length also in-
creases during the larval period from an average of 35% of
body length in preHexion larvae, to 36% in larvae under-
going flexion, to 37% in postflexion larvae. The eye is
moderate in size and averages 30% of the head length
during the larval period with no trend of relative increase.
Snout length averages 35% of the head length over the
letrval period.

The bases and blades of the pectoral fins are well dif-
ferentiated in the 2.8-mm specimen of Sparta (1942).
The short round blade portion of the pectoral fin is
another feature held in common with many species of
Sebastes. The paucity of specimens of H. dactylopterus
precluded staining for osteological study; however, the
size at formation of pectoral fin rays can be observed in
unstained specimens. Pectoral rays begin to appear when
the larvae are about 4.0 mm long and the last (lower-
most) rays have differentiated at about 8.0 mm. The
usual number of rays is 19. The pelvic fin buds appear in
larvae about 6.0 mm long. The rays begin differentiating
in 7-mm larvae, and the full complement of one spine
and five rays is present in the 8.6-mm specimen.

The hypural anlage of the caudal fin is apparent in 3-
mm larvae and the elements and principal caudal rays
begin to differentiate in 4-mm larvae. The full com-
plement of principal rays (8 + 7) is present in 7-mm lar-
vae. Notochord flexion occurs at a relatively large size
(6.0 to 8.0 mm) as in Sebastes and is another character
held in common by the two genera. The dorsal and anal
fins begin to form simultaneously in larvae about 6.0 mm
long, but the full complements of XII, 11 or 12 dorsal rays
and in, 5 anal rays are not present until 8.6 mm.

Larvae of H. dactylopterus have a distinctive pigment
pattern that is established in larvae less than 3.0 mm
long. On the head, a group of melanophores is present on
the lower jaw and above the brain. The dorsolateral sur-
faces of the gut are covered with a solid shield of pig-
ment which enlarges ventrally with continued develop-
ment. Several superficial melanophores are present on
the trunk just above the axillary region. The medial sur-
face of the pectoral fin base is solidly pigmented and fine
melanophores are located at two regions on the blade of
the fin, some near the distal margin and another group at

the basal region of the fin blade. A group of
melanophores is present at the ventral midline of the tail
just anterior to where the caudal fin will form. This uni-
que pattern of pigmentation remains essentially un-
changed throughout the entire larval period.

Distribution. — Adults of H. dactylopterus have a
complex distribution. Eschmeyer (1969) recognized two
Atlantic subspecies, H. d. dactylopterus and H. d. lahil-
lei, with the former composed of four separate pop-
ulations (northeastern Atlantic and Mediterranean, Gulf
of Guinea, South Africa, and northwestern Atlantic).
The subspecies H. d. lahillei is found off Uruguay and
Argentina. The 6.0- and 6.2-mm specimens in the series
are from Discovery station 714, off Uruguay, and thus are
larvae of H. d. lahillei. They are not distinguishable from
larvae of H. d. dactylopterus.

Sebastolobua Gill

Literature. — Pearcy (1962) described the floating egg
masses, the developing embryos, and the newly hatched
larvae oi Sebastolobus. The larvae, pelagic juveniles, arid
early demersal juveniles of S. altivelis and S. alascanus
are described and illustrated in Moser (1974). Larvae of
the other species in the genus, S. macrochir, of the north-
western Pacific, have not been described; however,
pelagic juveniles of this species are described and illus-
trated in Moser (1974).

Distinguishing features. — Early Sebastolobus lar-
vae (up to 6 mm) can be distinguished from those of all
other genera of eastern Pacific Scorpaenidae on the basis
of pigmentation. Sebastolobus larvae of this size range
are unique in having two large melanistic blotches about
midway along the tail, one at the dorsal midline and one
at the ventral midline. These are sometimes expanded to
form a solid band on the tail (Fig. 24). Early larvae of all
other eastern Pacific scorpaenid genera have a series of
melanophores along the ventral midline of the tail, and
in some species of Sebastes, an opposing row is present at
the dorsal midline. The large tail blotches of
Sebastolobus disappear in larvae between 4.2 and 6.4
mm. Soon after the loss of these large tail blotches the
larvae develop prominent crestlike parietal ridges that
terminate in double spines, the posterior (nuchal) spine
being longer and more prominent than the anterior
(parietal) spine (Fig. 24). Of the other eastern Pacific
scorpaenid genera, only the larvae of Scorpaenodes have
parietal ridges and spines like Sebastolobus. If two
spines are present on the parietal ridges of other genera,
the anterior spine is always longer and more prominent
than the posterior. Sebastolobus larvae may be dis-
tinguished from those of Scorpaenodes on the basis of a
melanistic shield, which covers the dorsolateral surface
of the gut in the former and is absent in the latter. Lar-
vae of Sebastolobus smaller than 10.0 mm could not be
identified to species. Larvae larger than this can be iden-
tified to species by a combination of characters that are

40

B

Figure 24.— Larvae of Sebastolobus spp. A. 3.0 mm; B. 3.5 mm; C. 5.2 mm; D. 5.7 mm; E. 6.2

mm; F. 7.7 mm.

41

Figure 25.— Developmental series of Sebattolobus altivelis. A. 11.2-mm larva; B. 26.g-mni pelagic juvenile; C. 53.5-mm pelagic

juvenile.

42

given in the following species descriptions of S. altivelis
and S. alascanus.

Sebastolobus altivelis Gilbert, Figure 25

Literature.— Larvae, pelagic juveniles, and early
demersal juveniles are described in Moser (1974).

Distinguishing features. — Larvae of Sebastolobus
hatch and are freed from their floating egg mass at about
2.6 mm in length. They have an elliptical yolk sac with a
posteriorly positioned oil droplet. When the larvae reach
approximately 3.5 mm, the yolk sac has been resorbed
and the jaws and feeding apparatus are well formed.
Flexion of the notochord begins in larvae about 6.0 mm
long and is completed in larvae about 7.5 mm (Table 26).
Transformation into the pelagic juvenile stage is in-
itiated within the size range of 14.0 to 20.0 mm (Table
27). All specimens larger than 20.0 mm have some juve-
nile pigmentation. The pelagic juvenile stage of S. altive-
lis is highly protracted and may last as long as 20 mo.
Transformation into demersal juveniles occurs within the
size range of about 42 to 56 mm.

Larvae larger than 10 mm may be differentiated to
species on the basis of body depth and pectoral fin shape.
Larvae and transforming specimens of S. altivelis are
more robust and deeper-bodied than those of S. alas-
canus. Body depth of specimens of S. altivelis in the 10-
to 20-mm size range is 36 to 45% of the body length (mean
of 41%), whereas in specimens of S. alascanus of compar-
able size the range is 33 to 39% with a mean of 35%. The
pelagic juveniles of S. altivelis are equally robust; body
depth averages 41% of body length in the 20- to 40-mm
size range but then decreases gradually to 34% in the
largest pelagic juveniles. Pelagic juveniles of S. alascanus

are less robust; body depth averages 34% with a range of
31 to 36%.

As in other scorpaenid genera, the pectoral fins provide
excellent characters for species separation. The pec-
torals are longer and have a deeper base in larvae and
pelagic juveniles of S. altivelis compared with S.
alascanus. In specimens of S. altivelis in the 10- to 20-
mm size range, the pectoral fin length averages 34% of
the body length (range of 29 to 39%). The pectorals reach
their maximum relative length in 20- to 40-mm pelagic
juveniles where they average 39% of the body length
(range of 35 to 41%). In pelagic juveniles 40 to 50 mm
long the average is 35% (range of 32 to 37%). In 10- to 27-
mm specimens of S. alascanus fin length averages 29% of
body length (range of 26 to 31%).

The two species can be separated by a combination of
meristic characters. Sebastolobus altivelis usually has
one or two more pectoral fin rays and one or two less ver-
tebrae than S. alascanus (Table 28).

The melanophore pattern of larvae of S. altivelis is in-
distinguishable from that of S. alascanus. At the begin-
ning of the larval stage (3.5 mm) there are large median
blotches opposite each other about midway back on the
tail. In some specimens, the blotches are expanded to
form a band. Also, melanophores cover the dorsolateral
surfaces of the posterior region of the gut. The melanis-
tic tail blotches are transitory; they are lost in some lar-
vae as small as 4.2 mm and one or both are absent in
most larvae between 5.0 and 6.0 mm in length. The dor-
sal blotch was absent in all larvae larger than 6.0 mm
and the ventral one was absent in all larger than 6.4 mm.
In contrast, the gut pigment is augmented throughout
the larval period, extending forward to the axillary region
and internally anterior to the cleithrum in larvae about
5.0 mm long. When the larvae reach 6.0 mm, the pig-
ment extends onto the ventral surface of the gut and dor-

Table 26. Measurements Cmm) of Sebastolobus spp. larvae. (Specimens between dashed lines are undergoing notochord flexion)

Standard

Snout-anus

Head

Snout

Eye

Body

Pectoral

Pectoral

Pelvic

Snout-

Snout-

lengtii

distance

length

length

diameter

depth

fin length

fin base depth

lln length

dorsal fin

anal fin

3.0

1.3

0.50

0.08

0.20

0.60

0.3

0.14

3.5

1.4

0.80

0.18

0.25

0.52

0.23

0.20

3.8

1.6

1.0

0.22

0.27

0.68

0.40

0.38

4.2

1.8

1.2

0.25

0.30

0.86

0.45

0.40

4.7

1.9

1.2

0.31

0.30

0.80

0.46

-

5.0

2.0

1.4

0.35

0.33

0.90

0.65

0.50

5.2

2.1

1.4

0.43

0.40

1.0

0.62

0.55

5.5

2.3

1.6

0.42

0.35

1.1

0.65

0.58

5.7

2.3

1.7

0.56

0.40

1.2

0.95

0.85

5.8

2.4

1.7

0.51

0.39

1.2

1.0

0.78

5.9
6.0

2.6
2.7

1.8
1.8

0.62
0.55

0.47
0.51

1.6

1.8

1.0
1.3

0.75

0.85

O.iT

2~ '

~3T4

6.2

2.7

1.9

0.70

0.48

1.6

1.4

0.90

0.20

-

-

6.4

2.7

1.8

0.60

0.45

1.5

1.2

0.85

0.13

-

-

6.7

3.1

1.9

0.82

0.55

1.8

1.4

1.0

0.15

3.9

3.9

6.8

2.8

2.0

0.82

0.52

1.8

1.4

1.1

0.17

-

3.2

6.9

3.3

2.2

0.80

0.58

2.0

1.5

1.2

0.25

2.7

4.1

7.1

3.5

2.5

0.82

0.71

2.4

1.8

1.2

0.80

2.5

4.2

_ ^3 _

_3.8_

2.5

0.95

0.76

2.4

1.8

1.4

0.85

2.7

4.4

7.7

4.2

2.7

1.1

0.75

2.8

2.1

1.5

0.85

2.9 ~

" "475 ~

7.8

4.3

3.1

1.1

0.82

2.6

2.1

1.4

1.2

3.2

4.8

8.3

4.4

3.1

1.1

0.79

2.6

2.0

1.4

1.0

3.1

4.9

8.6

4.8

3.2

1.2

0.92

3.1

2.4

1.7

1.4

3.5

5.3

8.9

5.2

3.3

1.2

1.0

3.1

2.5

1.7

1.4

3.8

5.4

9.2

5.2

3.5

1.2

1.0

3.2

2.7

1.5

1.5

3.8

5.6

9.6

5.4

3.5

1.2

1.1

3.7

2.8

1.8

1.7

4.1

5.8

43

Table 27. Measurements (mm) of ScbBslolobus nltivclis . (specimens below dashed line have
completed transformation Into pelagic Juvenile stage, j

StanAird

Snout-anus

Head

Snout

Eye

Dody

Pectoral

Pectoral

Pelvic

Snout-

Snout-

length

distance

length

length

diameter

depth

fin length

fin base depth

rin length

dorsal fin

anal fin

10.1

6.2

3.8

1.2

1.2

3.8

2.8

1.8

1.7

4.2

6.4

10.5

6.5

4.2

-

1.2

3.9

3.0

1.9

1.8

4.4

6.7

11.2

6.9

4.5

1.3

1.4

4.2

3.5

2.1

2.4

5.2

7.3

12.7

7.3

4.6

1.3

1.5

4.7

3.8

2.0

2.5

5.4

7.9

13.4

8.0

5.0

1.5

1.6

4.8

3.7

2.0

2.5

5.4

8.3

14.4

8.8

6.2

1.7

2.1

5.7

5.0

2.7

3.1

6.0

9.6

15.2

10.1

6.1

1.8

1.9

6.3

5.2

2.8

2.9

6.7

10.6

15.4

9.8

6.0

1.7

2.2

6.4

4.7

2.8

2.9

6.9

10.1

15.4

9.9

6.5

1.9

2.2

6.3

5.8

2.9

3.8

6.8

11.0

15.7

10.3

6.8

2.1

2.1

6.9

5.8

2.9

3.8

6.7

10.8

16.0

10.3

6.2

1.8

2.0

7.1

5.4

3.1

3.3

7.0

10.6

16.3

10.3

7.2

2.1

2.2

6.6

6.2

3.0

4.0

6.8

11.2

16.6

10.8

7.4

1.8

2.4

6.6

6.4

3.0

3.4

7.6

11.8

16.7

11.2

6.7

1.9

2.2

7.5

5.8

3.2

3.3

7.1

11.3

17.6

11.5

7.6

2.0

2.5

7.2

6.8

3.2

4.2

7.1

12.2

17.6

11.7

7.0

1.9

2.4

7.3

5.9

3.1

3.8

7.7

12.0

18.2

12.0

7.7

2.1

2.6

7.3

6.3

3.1

4.2

7.8

12.7

18.4

12.5

7.1

2.0

2.5

8.3

6.2

3.6

4.2

7.5

12.8

19.3

13.2

7.9

2.1

2.5

8.5

6.7

3.8

3.8

8.3

13.4

19.4

13.0

8.5

2.3

2.5

8.7

7.6

3.8

4.7

8.4

14.0

20.7

13.8

8.5

2.6

2.3

8.4

7.6

3.7

4.8

8.6

14.7

21.3
22.6

14.9
15.9

9.2
9.6

2.8

2.5

2.8
3.1

9.0
9.2

8.1

8.8

3.8
4.2

4.8
5.6

9.5
9.5

15.2
16.7

23.4

16.6

9.8

2.5

2.7

10.1

9.0

4.4

5.6

9.5

17.4

24.5

17.7

10.8

2.9

3.4

11.0

10.0

4.5

5.8

10.1

18.1

25.8

18.6

10.6

2.9

2.9

11.3

9.3

4.4

5.9

10.1

18.9

26.8

17.7

10.6

2.8

3.8

10.6

11.0

4.8

6.5

10.1

18.8

27.7

20.3

11.5

2.9

3.6

12.2

11.0

5.4

7.1

11.7

22.0

28.3

20.3

12.0

3.2

3.7

12.2

11.5

5.0

7.1

11.S

21.8

29.4

20.4

12.3

3.3

3.8

11.5

11.5

5.2

7.2

12.3

21.1

30.5

21.1

13.0

3.8

3.6

11.7

11.7

5.4

7.6

12.3

23.0

31.4

21.1

12.3

3.2

3.5

13.7

12.2

5.8

7.5

12.3

22.9

32.4

21.1

12.5

3.5

3.6

13.0

12.8

5.8

8.3

12.5

22.2

33.6

23.1

14.0

4.2

4.2

13.5

11.7

5.8

7.5

12.8

25.1

34.6

23.5

13.4

3.8

3.8

14.2

13.5

6.0

8.2

13.7

25.5

36.5

25.0

15.2

4.2

4.8

15.5

14.7

6.7

9.2

14.9

27.5

37.5

25.8

15.0

4.2

4.6

15.9

14.4

6.7

8.3

14.7

27.6

38.5

25.6

15.2

4.0

4.8

16.0

15.7

6.6

9.8

15.2

27.5

39.5

27.9

16.2

5.0

4.8

16.0

15.0

6.7

8.9

16.0

29.9

40.4

27.1

16.4

5.0

4.8

15.2

15.0

6.8

10.8

16.0

30.0

41.6

27.9

16.4

4.3

5.8

15.9

15.2

7.1

10.1

16.2

29.6

42.3

28.1

16.9

4.6

5.2

17.4

15.2

7.2

10.0

15.9

29.4

43.5

30.1

16.9

5.2

5.4

16.0

14.7

6.7

9.5

17.4

32.4

44.4

30.0

17.9

-

4.8

16.4

16.4

7.1

9.5

17.2

32.2

45.1

29.5

18.2

4.8

5.4

17.6

16.0

7.8

10.1

17.2

32.0

46.2

30.7

17.9

5.0

4.9

17.7

17.1

8.1

11.5

17.1

33.5

47.1

30.2

18.1

5.0

5.6

17.1

15.2

7.0

10.3

17.7

33.0

48.5

32.0

18.6

5.4

5.8

18.6

16.9

7.8

10.5

17.7

34.7

49.5

32.2

19.8

5.0

5.8

18.6

18.1

8.3

11.8

16.4

36.2

51.3

37.0

20.6

6.7

5.8

18.8

17.4

8.0

11.3

18.9

40.0

53.5

35.7

20.6

6.2

7.5

18.6

16.9

8.2

11.0

18.7

37.8

56.0

37.6

21.4

5.8

7.7

19.2

19.8

8.3

12.3

20.3

40.6

•42.0

26.8

16.6

4.6

5.6

12.0

12.7

6.0

9.3

16.4

29.0

•47.4

29.6

17.7

4.7

6.9

13.9

13.5

6.8

10.1

18.1

31.7

•48.1

31.4

19.1

5.2

7.3

14.4

15.2

7.0

11.5

17.9

33.5

•50.3

33.0

19.2

5.4

7.3

14.0

13.8

7.1

11.8

18.8

35.3

•51.0

33.0

19.5

4.7

7.2

15.0

16.2

7.5

11.2

18.7

35.2

•53.0

33.4

20.5

5.4

8.6

14.6

15.5

7.0

12.1

19.9

37.7

•54.0

35.4

19.7

5.9

7.3

15.5

17.7

7.8

12.6

20.6

38.4

•56.0

35.2

21.1

5.6

7.7

15.5

16.4

7.4

12.6

20.8

38.1

•57.5

37.2

21.4

5.6

8.0

15.3

16.7

8.1

12.5

21.1

40.3

•58.6

37.9

22.6

6.0

8.7

16.5

16.9

8.2

12.7

21.6

40.6

•60.0

38.0

22.0

6.0

8.4

16.2

15.1

8.1

12.8

21.5

41.6

•Benthic Juvenile.

44

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45

sally as deeply embedded pigment at the nape. With
continued development the melanophores form a solid
sheath on the peritoneum surrounding the gut.

Melanophores appear at the posterior margin of each
pectoral fin in some larvae as small as 4.0 mm. About
half the larvae examined between 4.0 and 5.0 mm have
this posterior margin of fine melanophores and the pig-
ment is present in all larvae in the 5.0- to 11.0-mm range.
The melanophores are then lost, and almost all larvae
larger than 11.0 mm have unpigmented fins.

Melanophores appear on the posterior lobes of the
brain in 5.2- to 7.0-mm larvae and are present in all lar-
vae larger than this. They also appear above the anterior
lobes of the brain in larvae between 7.0 and 9.0 mm in
length and in most larvae larger than 9.0 mm.

Pigmentation of the pelagic juveniles of the two species
is markedly different. It begins to appear in some
specimens of S. altivelis of the 14- to 20-mm size range.
On the head, patches of melanophores appear on the
opercle, cheek, snout, and jaws. In most specimens larger
than 22 mm, the patches are confluent, and the head is
generally dusky with darker areas at the opercle and
along the upper jaw.

A patch of melanophores appears superficially over
each side of the gut in specimens as small as 14 mm.
These patches expand to form a solid melanistic sheath
in some specimens of the 14- to 20-mm size range. The
posterior margin of the sheath is an arc running from the
vent to the nape and stands out sharply against the pig-
mentless region of the trunk posterior to the sheath. With
continued development the pigment sheath expands
posteriad and is a striking feature of the pelagic
juveniles. In the 20- to 30-mm size range the sheath ex-
tends posteriad to a vertical from the first or second anal
fin spines. In the 30- to 40-mm size range the sheath ex-
tends posteriad to the 2nd or 3rd anal spine in most
specimens and to the soft dorsal fin in some. In most
pelagic juveniles of the 40- to 50-mm size range, the
dusky sheath extends back to the soft dorsal and it does
so in all specimens of the 50- to 60-mm size range. When
the juveniles become benthic, the dusky sheath extends
posteriad to the caudal fin.

The fins become deeply and characteristically pig-
mented in juveniles of S. altivelis. The anterior portion of
the spinous dorsal fin becomes melanistic in specimens
as small as 18 mm. In juveniles of the 20- to 25-mm size
range, the anterior one-half to two-thirds of the fin is
melanistic. In the 25- to 30-mm range, three-fourths or
more of the fin is black. In pelagic juveniles larger than
this the pigment has spread onto the soft dorsal fin, and
covers both the soft dorsal and soft anal fins in benthic
juveniles.

The base of the pectoral fins begins to be covered with
melanophores in specimens as small as 14 mm. In
specimens as small as 18 mm the melanophores extend
onto the basal region of the rays. With further growth the
black basal zone enlarges posteriad and becomes a highly
characteristic feature of the pelagic juveniles. The pos-
terior margin of this zone contrasts sharply with the dis-
tal clear region of the fin. In juveniles of the 19- to 25-mm

size range, the width of the black basal zone averaged
20^"? of the fin length. Enlargement of this zone is shown
by the average relative widths for successive size ranges
(25 to 30 mm, 38^c; 30 to 35 mm, 46%; 35 to 40 mm, 54%;
40 to 45 mm, 65%; 45 to 50 mm, 72%; 50 to 55 mm, 76%).
Towards the end of the pelagic juvenile stage a pale
translucent layer covers the basal region of the pectoral
fin and gives the black zone the appearance of a band.

Distribution. — An extensive discussion of the dis-
tribution and abundance of larvae, juveniles, and adults
of S. altivelis is given in Moser (1974).

Sebastolobus alascanus Bean, Figure 26

Literature. — Larvae, pelagic juveniles, and early
demersal juveniles are described in Moser (1974).

Distinguishing features. — Larvae of Sebastolobus
smaller than 10 mm are not separable to species. Dis-
tinguishing features ol Sebastolobus larvae up to this size
are given in the generic account and in the section de-
scribing S. altivelis. Sebastolobus alascanus larvae longer
than 10 mm differ from those of S. altivelis in being less
robust and in having shorter pectoral fins with a nar-
rower base and fewer pectoral rays (Tables 29, 30). These
characters carry through into the pelagic juveniles of the
two species. The descriptive details for larvae and
pelagic juveniles of both species are given in the section
on S. altivelis.

In transforming species of S. alascanus, the first head
pigment to appear is a melanistic blotch on the posterior
region of the opercle. This gradually spreads anteriorally
onto the cheek and in late-stage pelagic juveniles the en-
tire head is speckled with melanophores. A blotch begins
to form over each side of the gut in larvae as small as 15
mm. These enlarge dorsad onto the spinous dorsal fin
and posteriad as an irregular mottled sheath that con-
trasts markedly with the solidly pigmented sheath of S.
altivelis juveniles. In the largest pelagic juveniles the
mottling on the dorsal fin and trunk extends posteriad to
a vertical from the vent. When the juveniles become ben-
thic the mottling spreads onto the remainder of the body
and median fins. Melanophores appear on the bases of
the pectoral fins in specimens of the 14- to 20-mm size
range. A faint band of melanophores appears on the rays
in some specimens of this size range but never becomes
highly developed and covers only the basal one-third of
the fin in the largest pelagic juveniles.

Distribution.— An extensive discussion of the dis-
tribution and abundance of S. alascanus larvae,
juveniles, and adults is given in Moser (1974).

Trachyscorpia Ginsburg

Literature.— None.

Distinguishing features. — No larvae are known and
the genus is known in the eastern Pacific from one adult

46

Figure 26.— Developmental series of Sebastolobug alascaims. A. 10.3-inm larva; B. 16.0-min transforming specimen; C. 25.3-

mm pelagic juvenile.

47

Table 29. Measurements (mm) of larvae and Juveniles of Sebnstolobus alascanus.

Standard

Snout-anus

Head

Snout

Eye

Body

Pectoral

Pectoral

Pelvic

Snout-

Snout

length

distance

length

length

diameter

depth

fin length

fin base depth

fin length

dorsal Qn

anal fin

10.3

6.2

4.2

1.5

1.2

4.0

3.2

1.6

1.9

4.3

6.8

11.2

7.4

5.0

1.8

1.3

4.1

3.2

1.7

2.1

5.0

7.7

11.7

7.2

5.0

-

1.3

3.9

3.1

1.8

2.1

5.1

7.7

12.5

7.7

4.7

1.7

1.5

4.5

3.6

1.8

2.4

5.4

8.3

13.2

7.3

5.0

1.6

1.8

4.6

3.9

1.9

2.7

5.8

8.2

14.0

8.1

5.7

1.8

2.1

4.9

4.1

1.9

2.5

6.2

8.8

14.4

8.4

5.6

1.4

1.8

5.2

4.2

2.1

2.6

5.7

9.1

15.2

9.2

6.2

1.8

2.0

5.2

4.4

2.1

2.9

6.3

10.0

16.0

9.3

6.2

2.0

2.0

5.2

4.8

2.2

2.9

7.1

10,0

16.4

9.8

6.7

1.8

2.5

5.7

4.7

2.3

3.2

7.1

10.5

16.9

10.0

6.8

1.9

2.3

5.9

4.8

2.4

3.1

7.3

10.8

17.6

10.6

6.7

2.1

2.2

6.1

4.9

2.3

3.5

7.1

11.3

18.6

12.7

-

-

2.8

6.8

5.4

2.8

3.8

8.5

13.2

19.8

12.0

7.7

2.0

2.8

7.1

5.9

2.9

4.0

8.3

13.0

20.0

12.0

8.1

2.2

2.9

7.1

6.4

2.8

4.2

8.4

13.0

20.3

12.8

8.2

1.8

3.1

7.2

6.2

7.8

4.0

8.1

13.5

21.0

13.5

8.6

1.9

3.1

7.5

6.5

3.0

4.6

8.1

14.4

21.4

12.8

8.3

2.4

2.9

7.1

6.2

2.8

3.8

8.6

14.4

21.9

12.8

8.5

2.1

3.1

7.1

6.2

2.8

4.2

8.8

14.9

22.3

14.2

9.6

-

2.8

7.9

6.5

3.2

4.4

9.1

15.4

22.9

14.4

9.8

2.6

3.2

7.5

6.7

3.1

4.6

9.3

15.5

23.2

14.7

9.5

2.5

2.9

7.9

7.2

3.0

5.0

9.6

16.0

23.4

14.9

9.0

2.2

3.3

8.0

7.1

3.2

5.0

9.3

16.4

24.0

14.4

9.8

2.2

3.2

8.0

6.8

3.3

5.0

9.8

15.9

24.3

15.5

10.0

2.8

3.2

8.6

7.3

3.3

5.0

9.8

16.9

25.1

15.9

10.0

3.0

3.3

8.4

7.0

3.4

5.0

10,1

17.2

25.5

16.0

10.1

2.8

3.5

8.8

8.2

3.7

5.5

10.1

17.6

26.4

17.9

10.6

3.2

3.5

9.0

8.0

3.7

5.7

11.3

19.1

■27.2

17.9

11.2

3.0

-

8.3

8.1

3.3

5.4

10.8

19.3

*22.5

13.0

8.7

2.1

3.2

7.2

6.3

2.9

4.6

9.6

14.7

•25.3

16.9

10.3

3.5

3.2

8.1

6.8

3.3

5.0

11.2

18.9

•37.8

22.0

12.7

2.9

5.0

10.1

9.6

4.2

8.2

13.5

24.3

•39.0

24.0

14.1

4.0

5.1

9.9

9.8

4.0

8.8

15.0

26.7

•40.8

25.1

14.8

3.8

5.3

10.0

9.7

3.8

8.8

15.3

27.3

•42.3

27.7

15.4

4.3

5.8

10.1

10.0

4.2

9,0

16.3

29.6

•43.6

26.3

15.4

4.1

5.8

9.5

10.6

4.4

9.4

16.1

29.5

•44.7

26.2

15.9

4.3

5.8

11.2

10.5

4.8

8.1

16.9

29.7

•46.2

28.1

16.2

4.4

6.4

10.8

10.8

4.7

9.9

16.7

31.2

•48.8

29.1

16.8

4.7

6.5

11.4

11.4

4.7

10.5

18.0

32.9

•50,3

29.4

17.4

4.8

7.0

12.4

12.2

5.0

10.6

18.0

33.0

•51.0

30.6

17.5

5.0

6.8

11.8

11.5

5.2

10,5

17.5

33.6

•59.2

36.6

20.6

5.4

8.4

13.0

13.5

5.6

12.1

20,8

40.0

•60.0

36.8

21.6

5.7

9.0

14.3

14.3

5.9

13.0

21,5

39.7

* Benthlc Juvenile.

specimen reported as Trachyscorpia sp. (Chirichigno
1974). The specimen was collected off Peru at lat.
13°57.2'S and long. 76°42'W in 580 to 600 m. Esch-
meyer (pers. commun.), who has examined the specimen,
finds no obvious difference between it and T. crista-
lata from the Atlantic. If Eschmeyer (1969) is correct
in the placement of Trachyscorpia near Sebastolobus
in the subfamily Sebastolobinae, its larvae can be ex-
pected to share some of the features of Sebastolobus
larvae.

Scorpaenodea Bleeker
Literature. — The literature contains no descriptions

or illustrations of the developmental stages of Scor-
paenodes.

Distinguishing features.— Scorpaenodes larvae dif-
fer from all other known Scorpaenidae except Sebas-
tolobus, in having prominent crestlike parietal ridges
that terminate in double spines, the posterior (nuchal)
spines being more prominent than the anterior (parietal)
spine; in larvae of other known scorpaenid genera, if two
spines are present on the parietal ridge, the second is
smaller and subjacent to the first. Scorpaenodes differs
markedly from Sebastolobus in melanophore pattern
and in degree of development at comparable sizes. Newly
hatched Scorpaenodes larvae have a series of about a

48

09

m

9

a

J3

a
o

I

•a

a
a

o

a

I

"O

o

c

5

CO

S

•o
o

K>

a
o

a

o
>1

1

u

>

OOOOOOOOOOOi-tOOOOOO
00 CO CO CO CQ CQCOCQCOCOCOCQCOCOCOCOCOOO

3. "

J3

K

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1 >■ 1 1 1 1 1 1 t t 1 1 1 1 1 1

Dorsal
fin
rays

o o o

1 1 1 i; 1 1 1 1 1 1 1 1 1 1 1 1 1 1

c
c

<

= 2

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

S3S5BSS535n3B53333

m 5'

OaiOC«MCOMCOrtCOCOCOC3MTj*'^MCO

co^^tau>u3iac-<ococDio(otot-toc>c<-

_ "1

33 -S

I-

o

c

CMC^C4CMCs]0)OieMCMCMiNCMC4CaC^N09C4

o

a

3

W«-tWCSlCgWNCaC^NCgCM(MCaMM<NC4

■=5 5

5

CMCMNCS|(N<N01lMCMCa(N<MM(N«CM«CM

2

NMMC>JCNlCaC4CVlWN<NC4C^OJ<McaNN

Branchlo-
Btegal
rays

5

c-t-t*t*c~r-c*t^c-t-t-t-t-c-c»c*fr^

i-i

c*t*r-t*t»fr-t-t*t*fr-t-t-t-c-t-e»t«-t\

Secondary

caudal

fin rays

t4

o
1

■^lococDooaociooaaOQOaoQOcoo^oo

o

CD

a

3
CQ

COCOCOt»aOO)aOOOOCDO)COGOO>00>0

Primary

caudal
fin rays

O

1

o

3

aogoaoGDaOQOaocoQOOOGOQOaoaoaoaDnao

u30t*»ocg'^c^TrtDt£'aoeo'«*'eoeoio«5rN

49

dozen melanophores along the ventral midline of the tail.
These soon become embedded and are not visible in lar-
vae larger than 4.0 mm. Newly hatched Sebastolobus lar-
vae lack this series of melanophores, but have two large
melanistic blotches about midway along the tail. Scor-
paenodes larvae lack the melanistic shield that covers
the dorsolateral surfaces of the gut in larvae of Sebas-
tolobus, Scorpaena, and Sebastes. In place of this, small
Scorpaenodes larvae have a deeply embedded blotch just
dorsal to the axilla. This later enlarges to cover the dor-
sal surface of the gas bladder. Head spine and caudal fin
formation occur at smaller sizes in Scorpaenodes com-
pared with Sebastolobus. In Scorpaenodes, numerous

head spines are present in larvae less than 3.0 mm long,
while Sebastolobus of that size still have their yolk and
do not obtain a comparative degree of spine develop-
ment until they are twice that size. In Scorpaenodes,
notochord flexion occurs within the size range of 4.0 to
5.5 mm whereas in Sebastolobus it occurs in the range of
6.0 to 7.5 mm.

Scorpaenodes xyris (Jordan and Gilbert),
Figures 27-29

Literature. — There are no previous descriptions or il-
lustrations of Scorpaenodes larvae.

Fitfure 27.— Larvae ot Scorpaenodes xyris. A. 2.7 mm; B. 3.2 mm; C. 4.0 mm; D. 4.8 mm.

50

Figure 28.— Larvae of Scorpaenodes xyria. A. 6.2 mm; B. 6.2 mm, dorsal view; C. 9.2 mm; D. 13.0 mm.

51

Figure 29.— Developmental series of ScorpaenodeM xyru. A. 12.5-inm truuforming specimen; B. 11.8-mm benthic

juvenile; C. 12.7-mm benthic juvenile.

52

Distinguishing features. — A single species of Scor-
paenodes. S. xyris, is known from the eastern Pacific.
Comparison of S. xyris larvae with those of congeners is
not possible since we have only S. xyris larvae in our col-
lections.

The smallest larvae of S. xyris in our collections are 1.8
mm in length. These specimens have already exhausted
their yolk supply, so it is likely that newly hatched lar-
vae of S. xyris are slightly smaller than 1.8 mm. Larvae
reach a maximum length of 14.0 mm and transfor-
mation into benthic juveniles occurs within a size range
of 11.0 to 14.0 mm.

Scorpaenodes xyris larvae are deep-bodied; body
depth increases gradually during the larval period. It
averages 28% of body length before notochord flexion,

Table 31. Measurements {mn) of larvae of Scorpaenodes xyris. {Specimens between dashed
lines are undergoing notochord flexion.)

Snout-

Pectoral

Pectoral

Pelvic

Snout-anal

Standard

anus

Head

Snout

Eye

Body

fin

fin base

fin

fin

length

distance

lenqth

lenqth

diameter

depth

length

depth

length

distance

2.7

1.2

0.60

0,20

0.26

0,78

0.27

0.21

3.2

1-5

1.0

0.25

0.32

0.61.

0.61.

0.51.

3.8

1.8

0.37

0.32

1.0

0.97

0-67

-4;4-

1.6

— -r:?—

1.2

■'r:«—

O.kS

0.37
-6:35—

1.2

■y.Y'

1.1

0.66

'"X.Y"

'-"urre—

U.2

2-0

1.2

0.50

0.38

1.2

1.2

0.71

0.03

U.2

2.0

1-3

0.1.6

0.35

1.2

1,3

0.75

0.03

2,1.

''i

2.0

1-3

0.50

0.39

1.2

1.5

0.83

0.12

3.1.

k.k

2.3

1.5

0.1.8

0.35

1.1.

1.1.

0.85

0.15

2.6

k.b

2.1.

1,5

0.55

0.51

1.6

1.6

0.83

0.21

2-5

it. 9

2.8

2.0

0.72

0.5't

1.8

1.8

0.91

0.1.0

2-9

S.2

2.8

1.9

0,66

0.60

1.9

1,9

1.0

0.50

2.5

--i:J-

3-2

2.1

0.76

0-55

2.0

2,2

1. 1

0.70

3.2

j-j—

■To""

"6:70"

■■■6:58—

■y.y

""TT"

■ — r.-?—

— 0-50-

■—3:2—-

S.8

3.6

2.1.

0.75

0.65

2.3

2.2

1.1

0.80

3-6

6.2

3-6

2.3

0.78

0.70

2.1.

2.3

1.2

1 .0

3-7

6.S

3.6

2.1.

0.83

0.75

2.1.

2.8

1.2

1 .1.

3.9

6.B

k.z

2.6

0,92

0-77

2.7

2.5

1 .2

1 . 1

1..2

7.0

4-3

2.8

0.90

0.86

2-7

2.9

1.3

1-2

l.,5

7.2

It. 5

2.8

0.60

0,82

2-9

2.6

1-3

1.2

I..7

7-5

1..2

2.5

0.61.

0.86

3.0

2.8

1.1.

1.3

't.5

7-7

1..B

3-1

1-1

0.90

2,8

2.6

1.5

1,1.

1..8

7-9

ii.ii

3-1

1 ,0

0.86

2.8

2.9

1 1.

1.2

1..8

8.2

5-1

2,9

092

0-93

3.2

3.3

1.5

1.8

5-3

8.6

5.2

3.2

1.0

1.0

3.3

3-2

1.1.

1.7

5.3

9.0

5.'t

3.3

1.0

1,0

3.5

3.9

1.7

1-7

5-5

9.2

5.0

3.5

1.2

1.0

3.1.

3.6

I.S

1.5

5.3

9-5

6.3

3.9

1 2

1.2

3.8

3.5

1.6

1.8

6.1.

9.6

5-6

3.6

1 . 1

1.1

3-6

I..3

1-7

1.7

6-1

10. 0

6.!

3.6

1 .1

1-2

I.. 1

l.,5

1.8

1.9

6.2

10.3

6.8

1..1

1.2

1-3

1..I

3.8

1.8

1.7

6.8

10.6

6.9

I..1

1.2

1.3

3.8

3.6

1,8

1-7

7,0

n.o

6.1-

I.-2

I-)

1 .ii

1..2

1..2

1.8

2-0

6-5

11.7

7-7

I.-6

1.3

l.t.

k.i

1..2

2.2

2.1.

7-9

12.2

7-3

'..3

1 .<*

1.5

5.0

5.2

2.2

2.6

7.8

13-0

8.6

l*.i

1.2

1.6

5.2

5.2

2.1

2.2

8.8

•It .0

6.8

3-9

1-0

1.3

3-8

3-9

1.9

2-1

7.3

'11.8

7.6

I..6

1.2

1.1.

U.2

1..I

2.0

2-3

6.1

•12.5

8-1

1..6

1-2

1.7

I., 6

I..9

2.2

2.1.

6.7

««12.5

6.2

I.. 8

I.I

1.8

1..1.

I..2

2.1

27

8.6

*Transfonning specimen.
**6enthic juvenile.

33% during flexion, and 39% after the completion of flex-
ion (Table 31). In the slightly more elongate trans-
forming specimens and newly transformed juveniles,
body depth is 36% of body length.

The gut is compact in small larvae of S. xyris. Snout-
anus distance undergoes a relative increase throughout
the larval period. It averages 46% of body length prior to
notochord flexion, 52% during flexion, 61% in post-
flexion larvae, and 64% in transforming specimens.

The large spiny head is a prominent feature of S. xyris
larvae. Head length increases from a mean of 30% of
body length in preflexion larvae to 34% in larvae under-
going notochord flexion, to 38% in postflexion larvae and
transforming specimens. The eyes are relatively small;
eye diameter averages 30% of the head length for the en-
tire larval period with no obvious trend of relative in-
crease or decrease. The snout length is relatively short in
newly hatched larvae (25% of head length), increases to a
relative maximum length during notochord flexion
(average of 36% of head length), and then gradually
decreases during the remainder of the larval period to
25% of head length.

The smallest larvae (1.8 mm) in our collection have
pectoral fins with the base and blade well differentiated.
The fins enlarge rapidly and are prominent fan-shaped
structures in larvae larger than 3.0 mm. Fin length
averages 22% of body length prior to notochord flexion,
34% during flexion, and 39% during the remainder of the
larval period. They shorten during transformation (aver-
age of 36%) and are about 35% in newly transformed
juveniles. The base of the pectoral fin is wide as in other
scorpaenine genera. It averages 18% of the body length
for the entire larval period. Ossification of the rays begins
in 4-mm larvae (Table 32). The full complement of 16 to
19 rays is present in larvae longer than 4.7 mm.

The anlage of the caudal fin is present in 2-mm lar-
vae. The hypurals begin to ossify in 5-mm larvae and the
full complements of 3 superior and 2 inferior elements are
ossified in 8-mm larvae. The principal caudal rays begin
to ossify in 4-mm larvae and the full complement of 8

Table 32. Meristics from cleared and stained larvae of Scorpaenodes xyris.

Length
(Mm)

Principal
caudal
fin rays

Procurrent
caudal
fin rays

Branchio-
stegal
rays

Pectoral
fin
rays

-

Hypural
elemente

G111
raVers
(right

arch)

Anal
fin
rays

Dorsal
fin
rays

Pelvic
fin
rays

Vertebrae

supe-
rior

Infe-
rior

supe-
rior

Infe-
rior

left

right

left

right

supe-
rior

Infe-
rior

upper
limb

lower
limb

left

right

;.2
11.6
5.2
5.8
6.0
6.7
7.2
7.9
8.2

S.5

10.3

II. 7

8
8
8
8
8
8
8
8
8

e

8

7

17
16

17
16
17
17
16
17
18
17
18
17
19

0

5
6
7
7
7
7
8
9
10
10
9

1 11-5
1 1 1-5
111-5
1 11-5
1 1 1-5
111-5
III-5
lll-S
1 1 1-5
1 1 1-5

xill-7
XI 1 1-9
xlll-9
XI 1 1-9
Xlll-9
Xlll-9
XI 1 1-9
Xlll-9
Xlll-9
Xlll-S

1-2

1-5

18

53

superior and 7 inferior rays is present in a 4.6-min larva.
The procurrent caudal rays begin to ossify in 5-mm lar-
vae and the full complements of 4 superior and 4 inferior
rays are present in 6-mm larvae.

The pelvic fin appears in 4-mm larvae and increases in
length to 13% of body length at the completion of noto-
chord flexion. Fin length averages 18% of body length for
the remainder of the larval period. The rays begin to
ossify in 4-mm larvae, and the full complement of 1 spi-
nous ray and 5 soft rays is present at 4.6 mm in length.

The dorsal and anal fins begin to develop simul-
taneously in 4-mm larvae. Ossification of the rays begins
in 5-mm larvae and the full complements of XIII, 9 (10)
dorsal rays and III, 5 anal rays are present before the lar-
vae reach 6.0 mm.

Pigmentation is sparse in S. xyris larvae. The smallest
larvae in the collection have a series of 12 to 18
melanophores along the ventral midline of the tail, from
the junction of the gut to the hypural anlage. They also
have a patch of melanophores along the ventral midline
of the gut and several melanophores on the terminal sec-
tion of the gut. Also, there is an embedded blotch just
dorsal to the axilla and the distal region of each pectoral
fin is covered with fine melanophores.

As development proceeds some of this original pig-
ment pattern is lost. The series on the ventral midline of
the tail is lost before the larvae reach 4.0 mm. The
melanophores at the ventral midline of the gut are lost in
larvae between 4.0 and 4.5 mm. The patch median to the
axillary region enlarges to form a shield over the gas
bladder and is visible throughout the larval period.

Just before transformation, a group of melanophores
appears on the opercle and preopercle. Transforming
specimens from bottom collections have a striking mot-
tled pattern on the head and body. In some, the distal
mtirgin of the pectoral fin is still melanistic.

Distribution. — Scorpaenodes xyris is a coastal warm-
water species. Adults have a latitudinal range from Peru
northward to San Clemente Island, Calif., and occur off
islands such as Guadalupe and the Galapagos (Miller
and Lea 1972). CalCOFI surveys show that the larvae oc-
cur throughout the lower two-thirds of the Gulf of
California. On the outer coast, S. xyris larvae were found
only as far north as line 117 (about lat. 28°N) off Punta
Eugenia. The planktonic collections of the E ASTRO PAC
expedition show that S. xyris larvae occur from lat. 20°N
to lat. 20°S along a coastal band (Fig. 30). Larvae oc-
curred in small numbers in both CalCOFI and the
eastern tropical Pacific hauls. The largest number taken
on any haul was seven, and three-fourths of the positive
hauls had a single larva of S. xyris.

Larvae were taken on all CalCOFI cruises in the Gulf
of California (Fig. 30). More larvae were taken in June
than in the other months (February, April, December),
thus suggesting a summer spawning peak in this region.
On EASTROPAC expeditions, the number of occur-
rences was about equal on the winter cruises (February-
April) and the summer cruises (July-September).

SO'Pl'iMMijMM

• •

III. I J.I 1,1.1.1,1, lilil.l.l.lil.M.lilil I.lilililil.l. 1, 1, 1,1, 1 H

Figure 30.— Stations at which larvae of Scorpaenodes xyria were
taken on CalCOFI cruises (triangles) during 1956 and 1957 and
EASTROPAC expeditions (dots) during 1%7 and l%g.

Scorpaena Linnaeus

Literature. — The early stages of two North Atlantic-
Mediterranean species of Scorpaena, S. porcus and S.
scrofa, were described by Raffaelle (1888) and Fage
(1918). More thorough descriptions of the egg, larval, and
juvenile stages of these two species and of the eggs and
larvae of S. notatus were given by Sparta (1941, 1942,
1956). The latter paper includes a color plate. The early
developmental stages of the sculpin, Scorpaena guttata,
of California and Baja California were described from
specimens cultured in aquaria (Barnhart 1932; David
1939; Orton 1955). These authors described the egg
masses, developing eggs, and larvae up to the stage of
yolk exhaustion (about 3.0 mm body length).

Distinguishing features. — Scorpaena larvae are easi-
ly distinguished from Sebastes and Sebastolobus by
myomere count, Scorpaena having 24, Sebastes having
26 or more, and Sebastolobus having 28 or more. Scor-
paena differs further from Sebastolobus in having a row
of melanophores along the ventral midline of the tail, as
opposed to a single dorsal and ventral blotch, and in hav-
ing nonbifurcate parietal ridges. This latter character
also separates Scorpaena from Scorpaenodes. Scorpaena
larvae develop a moderately long pair of parietal spines
that reach their greatest relative length at about the
completion of caudal fin formation. A small nuchal spine
forms subjacent to each parietal spine. Scorpaena may
be separated from Pontinus on the basis of gut pigmen-
tation. Scorpaena larvae develop a melanistic shield over
the dorsolateral surface of the gut, whereas Pontinus lar-
vae have a deeply embedded blotch above the axillary
region which, as in Scorpaenodes, enlarges to cover the

54

dorsal surface of the gas bladder. This character is par-
ticularly useful in separating small larvae of Scorpaena,
Pontinus, and Scorpaenodes that have not yet formed
head spines. Another melanistic character useful in
separating Scorpaena larvae from those of Scorpaenodes
and Pontinus is the presence of a ventral midline blotch
just anterior to the juncture of the cleithra in Scorpaena.
This is absent in the other five genera.

Scorpaena larvae have highly characteristic pectoral
fins. They are moderate in length, as opposed to the ex-
tremely elongate fins of Scorpaenodes, and are fan-
shaped in contrast to the aliform pectorals of Pontinus.
Pigmentation varies among the species of Scorpaena, but
the pectorals are usually heavily pigmented.

According to a recent revision, there are nine species of
Scorpaena in the eastern Pacific (Greenfield 1974). Scor-
paena larvae usually occur nearshore, thus are not com-
mon constituents of our plankton hauls. Our collections
contain sufficient numbers of larvae for the following
descriptions of S. guttata and another form which we
designate as Type A.

Scorpaena guttata Girard, Figure 31

Literature. — The three publications on the early
developmental stages of S. guttata listed in the
preceding section describe stages up to 3.0 mm. Later
larval stages of S. guttata have not been previously de-
scribed.

Distinguishing features. — The early stages of Scor-
paena guttata have been studied in detail (Barnhart
1932; David 1939; Orton 1955) and the following account
is a summary of these studies. The egg masses are spawn-
ed at about midnight and float to the surface. They are
bilobed gelatinous structures, each lobe measuring 16 to
26 cm in length. The matrix is about 2-mm thick and
within it the eggs are evenly spaced in a single layer. The
slightly elliptical eggs measure about 1.2 mm at the long
axis. The yolk mass is colorless and contains no oil
globule. The chorion is colorless, transparent, and un-
sculptured. There is no apparent perivitelline space. The
eggs hatch when freed from the matrix at an average
time of 3 days after fertilization in southern California
waters. The newly hatched young are 1.9 to 2.0 mm long,
have a large elliptical yolk sac, and have a voluminous
dorsal finfold that is inflated in appearance. The pec-
toral fin buds are small and inconspicuous. A patch of
dendritic melanophores covers the dorsal aspect of the
gut and similar melanophores are also below the gut.

At about 4 days after fertilization, the larvae are 2.5 to
2.7 mm long, have 22 to 24 myomeres, and have a row of
melanophores along the ventral midline of the tail. At
about 5 days after fertilization, the yolk is half utilized,
and at about 6 days the mouth is formed. At 7 to 8 days
the yolk is depleted, the jaws are functional, and the pec-
torals are fan-shaped and have one to several rows of
melanophores along their distal margins. The previously
undescribed later larval stages of S. guttata are
delineated below.

Early-stage larvae of S. guttata are deep-bodied,
become comparatively more slender during notochord
flexion, and thereafter become increasingly deep-bodied.
Body depth averages 369c of body length prior to
notochord flexion, 30% during flexion, and 40% fol-
lowing flexion (Table 33).

The compact gut increases gradually in relative length
during the larval period. Snout-anus length averages 49%
of body length prior to notochord flexion, 51% during
flexion, and 60% following flexion.

At the completion of yolk utilization the head of S.
guttata larvae is moderately large and increases in
relative size during later larval development. Head
length averages 30% of body length in preflexion larvae,
34% during flexion, and 38% following flexion. The eyes
are moderately small; eye diameter averages 32% of the
head length throughout the larval period with no ob-
vious trend of relative increase or decrease. Snout length
increases from an average of 31% of the head length in
preflexion larvae to 33% in larvae undergoing flexion,
then decreases gradually to 28% in our largest larva.

The pectoral fins are small and poorly differentiated in
2.0-mm larvae but growth and differentiation is rapid
and, when the larvae have reached 3.0 mm, the fins have
a well-differentiated base, a deep fan-shaped blade, and
have doubled in length to 15% of the body length. Fin
length increases to an average of 21% of the body length
at notochord flexion. Following notochord flexion, fin
length averages 22% of body length up to about 10 mm,
and then there is a considerable jump in relative fin
length to 29% in our largest larva (12.8 mm). The depth
of the fin base is slightly smaller than in other scorpae-
nine genera. It averages 15% of the body length over the
entire larval period. Fin rays begin to ossify in larvae
about 4.0 mm long and the full complement of 17 to 19
rays is present in 5-mm larvae.

The hypural anlagen of the caudal fin are apparent in
larvae as small as 2.5 mm. They begin to ossify in 5-mm
larvae and in the largest stained specimen (6.5 mm) the
large superior and inferior elements are well ossified. The

Measurements (itm) of larvae of Scorpaena guttata- (Specir
are undergoing notochord flexion.)

. between dashed lines

Standard
length

Snout-
anus
distance

Head
length

Snout
length

Eye
diameter

Body
depth

Pectoral

fin
length

Pectoral
fin base

depth

Pelvic

fin
length

Snout-anal

fin
distance

2.0

t.l

0.38

o,n

0.18

0.90

O.U

0-16

-

-

2.9

1.4

0.95

0.27

0.26

1.1

0.45

0.37

-

3.2

1.6

1.0

0.30

0.32

1.1

0.57

0.50

3.9

1.7

1.2

0.35

0.35

1,3

0.60

0.62

4.2

2.1

1.2

0.44

0.45

1.2

0.93

0.69

0.08

-

4.5

2.2

1.6

0.50

0.45

1.3

1.0

0.75

0.10

2.6

4.8

2.4

1.7

0.52

0,52

1.4

1-0

0.78

0.12

2.7

5.4

2.7

1.7

0.60

0,56

1.5

1.0

0.83

0.11

3.1

5.7

3.2

2.0

0.67

0.67

1.8

1.2

0.94

0.32

3.4

5.8

3.2

2.0

0.6B

0.68

2.1

1.2

1,0

0.55

3.3

6,0

3.5

2.2

0.76

0.72

2.2

1.4

1.0

0.60

3.6

6.5

3-7

2-3

0.80

0.78

2.4

1.5

1.1

0.73

3.7

8-0

5.t

3.2

I.l

1,1

3.5

1.8

1.3

1.6

5.2

9.2

5. 5

3,8

1.2

1.0

3.8

2.0

1.4

1.6

5.6

9-8

6.0

3,8

1.1

1.1

3.9

1.8

1.5

1.7

6.1

12-8

7.9

5,0

1.4

1.8

5.4

3.7

1.8

2.6

8.2

55

B

Figure 31.— Larvae of Scorpaena guttata. A. 2.1 mm; B. 3.1 mm; C. 6.2 mm; D. 12.8 mm.

56

full complement of 8 + 7 principal caudal rays is under-
going initial ossification in a larva 5.2 mm long. Second-
ary caudal rays begin ossifying in 5-mm larvae. In the
largest stained specimen (6.5 mm) there are 4 superior
and 4 inferior secondary elements ossifying.

The pelvic fins appear when the larvae are about 4.0
mm long and increase in relative length to 10% of the
body length at the completion of notochord flexion.
There is a gradual elongation of the fin during later lar-
val stages to a maximum of 20% of the body length in our
largest larva. Pelvic fin rays begin ossifying in 5-mm lar-
vae and the full complement of 1 spinous ray and 5 soft
rays is ossifying before the larvae reach 6.0 mm.

The dorsal and anal fins begin to form simultaneously
in larvae about 4.5 mm long. Ossification begins in 5-mm
larvae and the full complements of III, 5 or 6 anal rays
and XII, 8 to 10 dorsal rays is present in 6-mm larvae.

Larvae of S. guttata are characteristically pigmented.
Pigmentation of larvae less than 3.0 mm long has been
described by other investigators and is summarized
above. Notable changes occur in the initial pigment pat-
tern in larvae larger than 3.0 mm. Melanophores first ap-
pear on the posterior margin of the pectoral fin in early
larvae and when the larvae are 3.2 mm long, the distal
half of the fin is covered. At 4.0-mm length, the entire fin
is covered with melanophores as is the medial surface of
the fin base. As the rays develop, the pigment becomes
restricted to the membrane between the rays and
becomes strikingly heavy. In 6-mm larvae the distal mai-
gin of each fin begins to lose pigment and this clear zone
widens to about 20% of the length of the fin in the largest
larva. Gut pigment was originally on the dorsolateral sur-
face of the gut, but spreads ventrally to cover all sur-
faces of the gut when the larvae reach 4.0 mm. There is a
gradual reduction in the number of pigment spots in the
ventral tail series. In larvae about 3.0 mm long the num-
ber of melanophores in this series ranges from 2 to 7 with
a mean of 4. 15 ± 1.62 SD for 50 larvae. In larvae between
3.0 and 8.0 mm there is a general decline in this number
but the average number for the whole size range is
2.64 ± 1.12 for 11 larvae. Larvae longer than 9.0 mm had
no ventral tail melanophores.

When the larvae reach 4.0 mm, a small blotch of pig-
ment develops just anterior to the cleithral symphysis.
This remains throughout the larval period and is aug-
mented by several more melanophores in larvae larger
than 9.0 mm. Melanophores develop on the head region
in larvae of about 4.5 mm in length. A single melano-
phore is embedded at the nape and persists through the
remainder of the larval period. It is augmented by several
more melanophores in larvae 8.0 mm and longer. A pair
of melanophores appears above the optic lobes of the
brain in 4. 5-mm larvae and to these are added other
melanophores until the optic lobes are covered in larvae
larger than 9.0 mm. A pair of melanophores forms above
the olfactory lobes in larvae about 6.0 mm long and per-
sists throughout the remainder of the larval period.
Several melanophores form around the base of the pel-
vic fins in larvae about 6.0 mm long, but the fins them-
selves remain unpigmented until late in the larval

period. Melanophores begin to form in the epaxial
myosepta above the pectoral fin in some larvae as small
as 6.2 mm and cover an extensive area in larvae larger
than 10.0 mm. Melanophores appear on the cheek and
below the eye in 9-mm larvae and become more exten-
sive in larger larvae.

Distribution. — Adults of S. guttata are known from
Santa Cruz, Calif., southward to Uncle Sam Bank, Baja
California (about lat. 24°35'N). This species also occurs
at Guadalupe Island, Baja California, and in the
northern part of the Gulf of California (Miller and Lea
1972). Distribution of S. guttata larvae was analyzed for
the 1966 and 1969 CalCOFI cruises. Larvae were taken
from June to November in 1966 and from July to Sep-
tember in 1969. Spawning appears to peak in August, as
44% of the total occurrences and 67% of the larvae for
1966 were taken during this month (Table 34).

Table 34. Numbers of occurrences and standardized numbers of larvae of
Scorpaena guttata taken by CalCOFI in 1966.

CalCOFI

line

Latitude^

June

Ju

y

August

September

October

November ]

Occ.

No-

Occ.

No-

Occ.

No.

Occ.

Ho.

Occ,

No.

Occ.

No.

97

32-1;. 5'

1

.

.

100

Sl'-l?.?'

1

6

-

103

31 "07.0'

I

3

1

2

12

107

30''27.9'

2

6

110

29*'52.0'

2

113

29=24.2 ■

1

3

1

28

117

28=58.0'

3

16

6

104

15

1

6

118

28"18.S'

1

9

119

28°19.0'

1

120

ZB-SS.O"

4

12

4

52

1

3

133

27°26.2"

1

2

28

1

3

127

26°S7.5*

4

12

1

62

130

26?33.5'

1

1

133

26-08.5'

1

4

29

137

26" 36.1'

"

-

2

"

1

2

1

8

Latitude of most shoreward station.

Larvae of S. guttata occur in the southern portion of
the CalCOFI pattern, from about the Mexican border
south to line 137, just north of Magdalena Bay (Table
34). The greatest concentration of positive hauls was off
central Baja California in the area around Punta
Eugenia (Fig. 32).

Scorpaena Type A, Figvire 33

Literature. — This larval form of Scorpaena has not
been previously described or illustrated.

Distinguishing features. — Egg masses of Scorpaena
species other than S. guttata have not been reported in
the eastern Pacific and yolk-sac stages of Type A larvae
are not present in our collection. Type A larvae are
similar morphologically to those of S. guttata, but are
comparatively more slender prior to notochord flexion.
Body depth at this stage averages 27% of body length for
Type A larvae (Table 35) and 36% for S. guttata. Body
depth for Type A larvae undergoing notochord flexion
averages 29% of body length, and for postflexion stages
the mean is 38%. Comparative percentages for S. gut-
tata larvae are 30% and 40%.

57

' I ' ' '

1

1

J

-

K "^

--

^

\-

r BANC 1 SCO

-

\

^ \

\^

,SS,„o-

■

■

\u^

^fe /OIEGO /

■

oS rs "■'-■

-

-

\

'°^:P^q\\

-

\

\ ° /^

^ -

-

\^^^^

%,

-

V

-

-

-

-

1 1 1

1

1 . I

1

Figure 32.— Stations at which larvae of Scorpaena guttata were
talcen on CalCOFI cruises during 1966. Solid circles indicate stations
where number of larvae exceeded mean (7.4) for all positive stations.
Area of complete grid is outlined (see Ahlstrom 1961 for complete
grid).

Table 35. Measu

ements

tim) of larvae of Scorpaena Tvpe A.
are undergoing notocnord flexion.)

(Specimen

between

dashed line

Standard
length

Snout-
anus
distance

Head
length

Snout
length

Eye
diameter

Body
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin
distance

2.2

1.2

0.58

0.20

0.22

0.57

0.23

0.26

2.5

1.2

0.70

0.23

0.25

0.67

0.27

0.32

-

-

2.8

1.4

0.73

0.21

0.22

0.68

0.28

O.40

-

-

3.0

1.4

0.87

0.25

0.30

0.90

0.42

0.43

-

3.5

1.8

1.0

0.33

0.32

0.95

0.52

0.52

-

-

3.7

1.8

1.0

0.34

0.33

0.98

0.47

0.55

4.0

2.1

1.2

0.41

0.40

1.2

0.65

0.60

0.05

2.2

4.2

2.1

1.3

0.40

0.40

1.3

0.65

0.65

0.07

2.3

4.6

2.3

l.S

0.48

0.42

1.4

0.75

0.68

0.10

2.6

5.0

2.7

1.6

0.50

0.51

1.4

0.77

0.70

0.07

2.8

5.5

3.1

1.9

0.62

0.60

1.5

1.0

0.71

0.18

3.2

5.8

3.4

2.2

0.73

0.73

1.9

1.1

0.97

0.69

3.4

6.1

3.8

2.3

0.70

0.72

2.\

1.2

1.1

0.53

3.8

6.8

4.2

2,5

0.75

0.90

2.8

1.3

1.2

1.1

4.2

7.0

4.2

2.6

0.83

0.90

2.8

1.5

1.2

1.1

4.2

7.6

4.8

2.9

0.91

1.0

3.0

1.7

1-2

1,1

4.8

8.0

4.8

3.1

0.93

1.1

3.5

1.8

1.4

1.4

4.8

12.2

8.8

5.2

1.6

1.7

5.7

4.0

2.0

1.9

9.2

The gut is compact and increases in relative length
during the larval period. Snout-anus length averages 50%
of body length prior to notochord flexion, 53% during
flexion, and 61% following flexion. Head size is moderate
and increases relative to body length during develop-
ment. Head length averages 28% up to the beginning of
notochord flexion, 32% during flexion, and 38% fol-

lowing flexion. Snout length and eye diameter show no
change relative to head size and average 32% and 33% of
head length, respectively.

The fan-shaped pectoral fins are similar in size to
those of S. guttata. They increase in relative length from
an average of 12% of body length in preflexion larvae to
16% during flexion, and 21% following flexion. Depth of
the fin base shows a more gradual relative increase of
14%, 15%, and 17% for the three stages. Pectoral fin rays
begin to ossify in 3-mm larvae and the full complement
of 19 to 20 rays is present in 4-mm larvae. The pelvic fins
begin to form in 4-mm larvae and reach a maximum of
18%) of the body length in our largest larva. Ossification
is initiated in 4-mm larvae and the full complement of I,
5 rays is present at 6.0-mm length.

The hypural anlagen of the caudal fin appear in lar-
vae about 2.5 mm long. They begin to ossify when the
larvae reach 4.0 mm and the large superior and inferior
elements are well ossified at a length of 6.0 mm. Primary
caudal rays begin to ossify in 3-mm larvae and the full
complement of 8 -t- 7 rays is present at about 4.0-mm
length. Secondary caudal rays begin ossifying at about
4.0-mm length and there are 5 superior and 5 inferior
rays present in our largest stained specimen (6.0 mm).

The dorsal and anal fins begin to develop simul-
taneously in larvae about 4.0 mm long. Ossification is in-
itiated immediately thereafter and the larvae larger than
5.3 mm have III, 5 anal rays and XII, 8 dorsal rays.

The larvae of Scorpaena Type A differ from those of S.
guttata chiefly in the pattern of melanophores. As in ear-
ly larvae of S. guttata, those of S. Type A have
melanophores along the dorsolateral surfaces of the gut,
along the ventral midline of the tail, and on the pectoral
fins. The particular arrangement of melanophores on the
tail and pectoral fins is distinctly different in the two
forms and is a useful means of separating them. In Type
A larvae the ventral midline series extends along the en-
tire length of the tail from above the anus posteriad to
the hypural anlage of the caudal fin. In larvae of S. gut-
tata, the series is confined to the posterior half of the tail.
Accordingly, there are more melanophores in this series
in Type A; the range for preflexion larvae is 8 to 14 and
for larvae undergoing flexion is 4 to 8. Ventral tail
melanophores are not present in postflexion larvae. The
melanophore pattern of the pectoral fin is similar in lar-
vae of S. Type A and S. guttata up to about 3.0 mm in
length. At that size, a region of darker pigmentation
develops at the ventral aspect of the fin in Type A. This
area of darker pigment remains detectable from the sur-
rounding pigment in larvae up to about 6.0 mm in
length. This pigment does not develop in larvae of S. gut-
tata. As in S. guttata, pigment begins to develop on the
medial surface of the fin base at about 3.0 mm in length
and intensifies during the remaining larval period. In
Type A larvae of about 4.0-mm length the dorsal and up-
per posterior margins of the fin begin to lose
melanophores. This clearing becomes more extensive
with further development and in the largest larvae the
membranes of the most dorsal three rays and the pos-
terior one-third of the fin are clear. In S. guttata only a

58

Figure 33.— Larvae o( Scorpaena Type A. A. 2.5 mm; B. 3.5 mm; C. 4.6 mm; D. 8.0 mm.

59

narrow portion of the distal region of the fin becomes
clear in mid- and late-stage larvae.

Other pigmentation is not substantially different from
that of S. guttata and other known Scorpaena larvae. A
melanistic blotch appears just anterior to the junction of
the cleithra at about 3.0-mm length and remains
throughout the larval period. Melanophores form pos-
teriorly on the brain when the larvae are about 4.0 mm
long. At about 5.5 mm, one or two melanophores appear
above the olfactory lobes. In 7-mm larvae additional
melanophores form above the optic lobes and at 8.0 mm
the brain is covered. At 4.0-mm length an embedded
melanophore forms at the nape and remains throughout
the larval period. Myoseptal pigment is present in the
epaxial musculature above the pectoral fin in the 8.0-mm
larva.

that station the mean number of larvae for the other 25
stations in the Gulf was 12. The species that produces
larvae of Scorpaena Type A is a summer spawner since
no larvae were taken on the December, February, or
April cruises in the Gulf during 1956 and 1957.

Larvae of Scorpaena Type A occurred at two stations
of the EASTROPAC expedition in the vicinity of Panama
Bay and at two stations of the Scripps Tuna Ocean-
ography Cruise 58-1, one near Panama Bay and one off
the Tres Marias Islands (lat. 21°30'N, long. 106°30'W).
The low incidence of these larvae on these wide-ranging
expeditions probably reflects the near-shore nature of
spawning in the species producing the larvae.

Pontinus Poey

Distribution. — Most of our larvae of Scorpaena Type
A were collected on the CalCOFI cruise in the Gulf of
California during June 1957. They occurred at 24 stations
distributed throughout the entire Gulf (Fig. 34). They oc-
curred at two stations of the CalCOFI cruise during
August 1957, however, only a small portion of the plank-
ton from that cruise was sorted and the larvae may have
occurred more extensively on that cruise. Except for one
station, where the standardized number of larvae was
143, the larvae occurred in small numbers. Eliminating

Figure 34.— Stations at which lar-
vae of Scorpaena Type A were
taken on CalCOFI cruises in the
Gulf of California during 1956 and
1957.

Literature. — The literature contains no descriptions
or illustrations of the developmental stages of Pontinus.

The taxonomy of Pontinus in the eastern Pacific is
poorly known. At present there are five recognized and
six nominal species, however, unpublished information
of W. Eschmeyer (Calif. Acad. Sci.) indicates that the
species complement of eastern Pacific Pontinus is more
than double the presently recognized number. In fact, a
previously undescribed but abundant species was dis-
covered while examining museum specimens during the
present study. Unfortunately, the characters that
separate this host of species are mostly not the kind that
may be traced backward into larval series.

Distinguishing features. — Larvae of Pontinus can be
separated from those of other eastern Pacific genera by a
combination of characters. As in larvae of Scorpaenodes
and Ectreposebastes. Pontinus larvae lack the melanis-
tic shield that covers the dorsolateral surfaces of the gut
in Sebastes, Sebastolobus, and Scorpaena. In place of
this, larvae of Pontinus, Scorpaenodes, and Ec-
treposebastes have a deeply embedded medial blotch
just dorsal to the axilla. This blotch later enlarges to
cover the dorsal surface of the gas bladder. Pontinus lar-
vae differ from Scorpaenodes in several characters. In
Pontinus, a single elongate spine develops at the ter-
minus of each parietal ridge. In midstage larvae a small
subjacent nuchal spine develops posterior to each
parietal spine and remains inconspicuous throughout the
larval period. This contrasts greatly with the bifurcate
terminus of the parietal ridge in Scorpaenodes. The pec-
toral fins of the two genera differ in shape, relative
length, and pigment pattern. They are aliform in Pon-
tinus and fan -shaped in Scorpaenodes. The pectorals are
shorter in Pontinus; pectoral fin length averages 31% of
the body length in postflexion larvae of Type A and 27%
in Pontinus Type B; mean length was 39% in postflexion
larvae of S. xyris. The pectoral fin base is also narrower
in Pontinus; the depth of the fin base averages 14% of the
body length in postflexion larvae of Type A and 16% in
Pontinus Type B. Fin base depth averaged 18% in post-
flexion S. xyris larvae. Pigmentation of the pectoral fins
is limited to the distal margin of the pectoral fin in Scor-

60

paenodes whereas in Pontinus the pattern of pigmen-
tation is more complex and develops through sequential
stages. Larvae of Ectreposebastes are highly distinctive
and may be separated from those of Pontinus on the
basis of body depth and pectoral fin length. Body depth
averages 46% of body length for postflexion larvae of Ec-
treposebastes and 39^'c for postflexion larvae of Pontinus
Type A. The pectoral fin extends to the base of the
caudal fin in Ectreposebastes and extends only to the
midregion of the anal fin in Pontinus. Also, pigmen-
tation of the pectoral fin is markedly different as evident
in the illustrations of larvae of the two genera.

Despite the large species complement of Pontinus, we
can distinguish only three larval forms in our collection.
We have a complete larval series of the abundant form,
here called Pontinus Type A. It ties in with the juveniles
of a common but undescribed species of Pontinus dis-
covered in this study. A second, much less abundant
form, called Pontinus Type B, is distinguishable down to
a length of about 5.0 mm. Its largest pelagic juveniles
have a pigment pattern similar to the smallest known
benthic juveniles of Pontinus sierra (Gilbert) and it is
likely that Pontinus Type B is P. sierra. A third form,
here called Pontinus Type C, is known from a few small
larvae and is presently not identifiable.

Since larvae of Pontinus have not been previously de-
scribed, a detailed description of Pontinus Type A is
given below along with a brief description of Type B.

Pontinus Type A, Figure 35
Literature. — None .

Distinguishing features. — The smallest larva
available to us is a specimen 2.3 mm long that has no
yolk sac. When the larvae reach about 15 mm, pigment
saddles begin to form dorsally on the trunk. Although
there is no associated abrupt morphological change we
have designated this as the beginning of the pelagic
juvenile stage. This stage is protracted in Pontinus Type
A, our largest pelagic specimen being 27.4 mm long. Ap-
parently, there is a large size range over which Pontinus
Type A pelagic juveniles may become demersal since
bottom-caught specimens as small as 17.2 mm are pres-
ent in our collections.

Larvae of Pontinus Type A are deep-bodied. Body
depth increases from a minimum of 25% of the body
length in the smallest larva to 33% at notochord flexion
(Table 36). Body depth averages 39% in postflexion lar-
vae and pelagic juveniles. In small benthic juveniles,
average body depth is reduced slightly to 36% of the body
length.

The gut is compact in small larvae of Pontinus Type A
and undergoes a relative lengthening throughout the lar-
val period. Snout-anus distance averages 48% of the body
length before notochord flexion, 54% during flexion, and
6Kt following flexion. In pelagic and small benthic
juveniles, snout-anus length averages 64% of the body
length.

Table 35. Measurements (im) of larvae of Pontinus Type A. (Spectmens between dashed lines
are undergoing notochord flexion.)

Snout -

Pectoral

Pectoral

Pelvic

Snout-anal

standard

anus

Head

Snout

Eye

Body

fin

fin base

fin

fin

length

distance

length

length

diameter

depth

length

depth

length

distance

2.3

1.0

0.67

0.20

0.24

0.58

0.41

0.36

2.7

1.2

0.70

0.19

0.24

0.73

0.49

0.40

3.0

1.6

1.0

0.30

0.31

0.96

0.55

0.45

..iA..

....UZ...

..1.5...

..Q.H..

...0.30...

.-I.O..

...Q.65..

0.50

4.1

2.2

1.3

0.52

0.40

1.4

1.1

""0'."6"5" —

-o:fo""

^4

...4.6..

....2.5...

..1.5...

..(1.6J..

...(1.16...

..I.5..

.. 1.2

0.70

0.24

2.8

5.2

3.0

2.0

0.70

0.62

2.0

■i'r4'

..........

o:84

3:2

5.5

3.5

2.4

0.95

0.56

2.2

1.6

0.86

0.73

3.6

5.9

3.8

2.6

0.90

0.56

2.4

1.8

0.86

1.0

3.8

6.4

3.8

2.5

0.85

0.65

2.6

1.8

1.0

1.2

3.8

6.7

4.0

2.6

0.95

0.68

2.6

2.0

1.0

1.3

4.2

7.1

4.1

2.7

1.0

0.72

2.7

2.1

1,1

1.2

4.2

7.5

4.4

2.9

1.1

0.75

2.8

2.2

1.1

1.4

1.5

6.0

4.8

3.1

1.0

0.84

3.2

2.4

1.1

1.5

5.0

8.6

5.1

3.4

1.0

0.91

3.4

2.7

1.2

i.e

5.5

9.2

5.5

3.5

1.0

0.95

3.6

2.8

1.2

1.8

5.7

9.8

6.0

4.2

1.2

1.0

3.7

2.8

1.2

2.0

6.2

10.6

6.3

4.2

1.2

1.2

3.8

3.6

1.5

2.5

6.5

11.2

7.3

4.8

1.6

1.2

4.7

3,9

1.6

2.8

7.5

12.2

7.5

4.8

1.4

1.5

4.4

4.1

1.6

2.6

6.1

13.2

8.1

5.2

1.5

1.5

5.1

4.2

1.8

2.9

8.7

14.4

8.8

5.8

1.7

1,8

5.3

4.3

1.9

3.3

9.5

*I5.0

9.7

6.0

1.8

1.8

6.0

5.3

2.0

3.6

10.1

*16.4

10.0

6.2

1.8

1,8

6.5

5.4

2.2

3.3

10.5

•17.7

11.2

7.3

2,1

1,9

7.2

6.1

2.4

3.9

11.7

•19.6

12.7

7.8

2.1

2.1

8.0

6.7

2.6

4.2

13.4

•21.3

13.7

8.8

2.7

2.3

6.0

6.7

2.7

4.3

14.4

•23.0

14.7

9.6

2.7

2.6

6.3

7.6

2.8

4.6

15.5

•25.7

16.4

10.1

3.2

2.9

9.5

7.9

3.2

4.8

17.2

•27.4

17.9

11.7

3.2

2.6

9.8

8.8

3.3

5.1

19.3

"17.2

10.6

6.9

1.8

1.9

6.2

5.9

2.2

3.8

11.7

••18.9

12.2

7.5

2.1

2.2

6.7

5.7

2.4

4.2

12.8

••20.8

13.0

8.2

2.2

2.4

7.4

6.8

2.6

5.0

14.0

•*24 5

16.0

9.8

2.9

2.7

8.8

7.5

3.0

5.?

17.4

•Pelagic juvenile.
•■Benthic juveni te.

The head is large in Pontinus Type A larvae. Head
length increases from a mean of 30% of the body length in
preflexion larvae to 32% in larvae undergoing notochord
flexion, and averages 40% in postflexion larvae and in
pelagic and small benthic juveniles. The moderately
small eyes undergo a relative diminution during the lar-
val period. Eye diameter averages 32% of head length
before and during notochord flexion and is reduced to a
mean of 27% in postflexion larvae and pelagic juveniles.
Eye diameter increases slightly to a mean of 28.5% of
head length in newly transformed benthic juveniles.
Snout length increases from a mean of 32% of head
length in preflexion larvae to a mean of 41% in larvae
undergoing flexion. Thereafter the mean decreases to
33% in postflexion larvae and to 29% in pelagic juve-
niles.

The pectoral fins are prominent in the smallest larvae
in our collection. At this stage they have a slight aliform
shape; however, during later larval stages they become
distinctly aliform. Fin length increases from an average
of 20% of the body length in preflexion larvae to 26% in
larvae undergoing notochord flexion, and to 32% in post-
flexion larvae and pelagic and small benthic juveniles.
The depth of the pectoral fin base is not as great as in
other eastern Pacific scorpaenine genera but is greater
than in sebastine genera. It decreases from an average of
16% of the body length before and during notochord flex-
ion, to 14% in postflexion larvae and 13% in pelagic and
early benthic juveniles. Sequence of ossification of fin
elements was not analyzed because only a few larger lar-
vae could be properly stained. Apparently calcium in the
larvae leached out in preservative. Pectoral ray counts
for nine stained specimens ranged from 17 to 19.

The dorsal, anal, and pelvic fins begin to develop in
larvae about 4.0 mm long. The counts for the nine stain-
ed specimens were D. XII, 9; A. Ill, 5; P. I, 5. Hypural an-
lagen of the caudal fin are visible in the smallest larvae;

61

B

Figure 35.-Developmental series of Pontinua Type A. A. 2.3-mm larva; B. 4.1-mm larva: C. 8.0-mm larva; D. 24.5-

mm benthic juvenile.

62

the adult complement is 3 + 2 hypurals. The stained lar-
vae had 8 + 7 principal rays and 6 + 6-7 procurrent rays.
All stained larvae had 24 vertebrae. Gill rakers ranged
from 5 to 6 on the upper limb and 8 to 12 on the lower
limb of the first arch. There were 7 branchiostegal rays.

Early larvae of Pontinus Type A are sparsely pig-
mented. The initial melanophore pattern is similar to
that of Scorpaenodes. A series of 11 to 16 melanophores
(mean of 13 ± 1.3 SD for 14 larvae counted) is present
along the ventral midline of the tail. The series extends
from the anus posteriad to the hypural anlage of the
caudal fin. A wider space separates the ultimate
melanophore from the others. The number of
melanophores and their spacing is similar in Scor-
paenodes xyris except that in that species the series ex-
tends farther anteriad to the point of divergence of the
terminal section of the gut. In Pontinus Type A larvae
the ventral tail melanophores begin to become em-
bedded during notochord flexion and are not visible in
larvae larger than 6.0 mm.

Other pigment present initially is a medial blotch
anterodorsad to the axillary region. With further
development this expands to form a shield over the gas
bladder. It becomes progressively less visible with
thickening of the trunk musculature. Similar pigment is
present in Scorpaenodes and Ectreposebastes.

Pigment is present initially on the gut as a few faint
melanophores along the dorsal surface of the preter-
minal section of the gut and as a streak along the ventral
midline, with heaviest concentration below the liver.
Also, fine melanophores are present along the edge of the
finfold just anterior and posterior to the anus. These
areas of pigmentation persist in larvae up to about 5.5
mm in length and are present thereafter. When the lar-
vae reach about 10.0 mm in length, large melanophores
begin to appear internally on the perivisceral membrane
and, with further development, eventually cover the
entire membrane.

The other pigmentation present initially is on the pec-
toral fins. Minute melanophores cover the entire blade of
the fin and are present at the margin of the medial sur-
face of the fin base. This contrasts with the pattern in
early larvae of Scorpaenodes which have, at most, the
distal half of the fin blade covered with melanophores
and which have none on the fin base. The pattern of pig-
mentation on the pectorals of Pontinus Type A goes
through a series of distinct changes as larval develop-
ment proceeds. Towards the end of notochord flexion the
melanophores disappear from the fin base and begin to
recede from the basal region of the fin blade. The basal
clear area continues to enlarge and, at maximum reces-
sion in larvae of about 10 mm in length, pigment is
reduced to the distal margin. At about 10.0 mm the dis-
tal band begins to expand and becomes progressively
wider during the remainder of the larval period and
pelagic juvenile phase. At the end of the pelagic juvenile
stage only a small portion of the fin remains unpig-
mented. Newly transformed benthic juveniles have a
mottled pattern on the fins with a prominent dark blotch
near the base.

In larvae about 6.5 mm long, a melanistic blotch ap-
pears on the membrane between the 3rd and 4th spinous
rays of the dorsal fin. Additional blotches appear an-
terior and posterior to this as development proceeds, and
larvae larger than 7.0 mm have a series of blotches from
the 2nd and 3rd interradial membrane to the 5th or 6th
membrane. In pelagic juveniles the entire membrane of
the spinous dorsal fin becomes pigmented; however, a
much darker blotch is present between the 6th and 11th
spines. This dark blotch persists in benthic juveniles.

Melanophores appear on the brain in 10-mm larvae,
initially with a single pigment spot above each optic
lobe. Others form rapidly and the entire dorsal surface of
the brain is covered in larvae larger than 13.0 mm.

Saddles of melanistic pigment begin to form on each
side of the dorsal midline in 13-mm larvae, one between
the 1st and 2nd dorsal spines, and the other between the
5th and 7th spines. These enlarge to include the regions
between the 1st and 3rd dorsal spines and the 5th and
10th spines in larvae larger than 15.0 mm. Two other pig-
ment saddles appear in pelagic juveniles, one below the
soft dorsal fin and the other at the caudal peduncle. In
the largest pelagic juveniles, the anterior pigment sad-
dle covers the nape, the large one posterior to it bifur-
cates and extends ventrad to the horizontal septum, the
one below the soft dorsal extends ventrad to the base of
the anal fin, and the one at the peduncle is a weak band.
These markings break up into a more complex pattern in
benthic juveniles.

Pigmentation appears on other regions in pelagic
juveniles. At about 15.0-mm length, a blotch forms at the
midregion of each pelvic fin and the entire membrane of
the fin is pigmented at about 20.0-mm length. When
pelagic juveniles reach about 25-mm length, a blotch ap-
pears at the midregion of the soft dorsal and the soft anal
fin. These appear as extensions of the prominent trunk
band in benthic juveniles.

The head becomes pigmented in pelagic juveniles first
on the opercle and on the region above the opercle. At the
end of the pelagic phase, the entire lateral aspect of the
head is mottled.

Distribution. — The midwater trawl and plankton col-
lections of CalCOFI, EASTROPAC, and the STOR
group show the geographic distribution of larvae and
pelagic juveniles of Pontinus Type A (Fig. 36). CalCOFI
plankton collections show that larvae and pelagic
juveniles of Pontinus Type A occur in the Gulf of Califor-
nia up to about lat. 28°N. On the outer coast of Baja
California, pelagic juveniles of Pontinus Type A have
been taken as far north as Punta Eugenia (about lat.
28°N). Collections of the EASTROPAC expedition and
STOR show that larvae and pelagic juveniles occur
southward to the northern coast of Peru (about lat. 5°S).
It has an extensive distribution in the Gulf of Panama
and westward to the Galapagos Islands. In general, its
distributional pattern is similar to that of Scorpaenodes
xyris except that Pontinus Type A is more abundant in
offshore regions along the tropical American coast. Lar-
vae occurred in small numbers in both CalCOFI and

63

JO-U|l|l|l|l|l|IJI|

lii.l^i^i.i.i.l I. Li. 1,1,1,1,1, 1. 1,1, 1, 1, 1, 1,1,1

Figure 36. — Stations at which larvae and pelade juveniles of Pon-
tirms Type A were collected on CalCOFI cruises (triangles) during
1956 and 1957, EASTROPAC expeditions (closed circles) during 1967
and 1968, and Scripps Tuna Oceanography cruises (open circles) dur-
ing 1958 and 1959. Open square represents a station from the Scripps
Institution of Oceanography fish collection and closed square a sta-
tion from the University of California, Los Angeles, flsh collection.

EASTROPAC plankton tows. The mean number of
specimens for all plankton tows was 3.4 ± 6.36 SD with a
range of 1 to 34; 50% of all plankton tows had only a
single specimen.

In the tropical waters surveyed during the EAS-
TROPAC expedition, the number of occurrences was
about equal on the winter cruises (February to April) and
the summer cruises (July to September).

Pontinus Type B, Figure 37

Literature. — None.

Distinguishing features. — The smallest larva
available is a 5.0-mm specimen undergoing notochord
flexion. Dorsal pigment saddles are already beginning to
form at this size, and well-formed saddles are present in
specimens as small as 10 mm. In Pontinus Type A the
pigment saddles begin to form at about 15 mm, a size
chosen for convenience as the beginning of the pelagic
juvenile stage. Since pigment saddles begin to form in
midstage larvae of Pontinus Type B it is more ap-
propriate to consider the appearance of scales at about 10
mm as the event marking the beginning of the pelagic
juvenile stage. The largest pelagic juvenile in our collec-
tion is 23.1 mm. The size at transformation into benthic
juveniles is not known since the smallest benthic juvenile
of Pontinus Type B in our collection is 29.0 mm.

The smallest larva in our collection, 5.0 mm long, has
almost completed notochord flexion indicating that

notochord flexion is probably initiated in 4-mm larvae.
At 6.0 mm, notochord flexion is completed. Notochord
flexion occurs at a smaller size in larvae o{ Pontinus Type
A and is completed when the larvae reach about 5.0-mm
length.

Larvae of Pontinus Type B are deeper-bodied than
those of Pontinus Type A. Body depth at the pectoral fin
base in Pontinus Type B larvae undergoing notochord
flexion is 45 to 46'^"c of the body length (Table 37). In post-
flexion larvae the mean is 43%, and in pelagic juveniles it
is 41% . The more slender-bodied larvae oi Pontinus Type
A show an opposite trend. Body depth increases from a
mean of 33% of the body length in larvae undergoing
notochord flexion to a mean of 39% for postflexion larvae
and pelagic juveniles.

The compact gut is already masked by trunk mus-
culature in the smallest larva in our collection. Snout-
anus length is relatively longer in Pontinus Type B than
in Pontinus Type A. It averages 64% of the body length in
Pontinus Type B larvae undergoing notochord flexion
and 67% in postflexion larvae and pelagic juveniles while
in comparable stages of Pontinus Type A the means are
54% and 62%.

The head is larger in Pontinus Type B than in Pon-
tinus Type A. Head length in the former averages 45% of
the body length during notochord flexion and in post-
flexion larvae and 42% in pelagic juveniles. Relative head
length shows an opposite trend in Pontinus Type A,
averaging 32% in larvae undergoing flexion and 40% in
postflexion larvae and pelagic juveniles. The eye is
relatively larger in Pontinus Type B compared with Pon-
tinus Type A; eye diameter averages 32% of head length
for larvae and pelagic juveniles of the former and 27% in
comparable stages of Pontinus Type A. Snout length is
relatively shorter in Pontinus Type B compared with
Pontinus Type A. In the former it averages 30% of the
head length in larvae undergoing notochord flexion, 28%
in postflexion larvae, and 24% in pelagic juveniles. At
comparative stages of Pontinus Type A it averages 41%,
33%, and 29% of the head length.

Pectoral fin length averages 33% of the body length in
Pontinus Type B larvae undergoing notochord flexion
and 27% in postflexion larvae and pelagic juveniles. In
comparable stages of Pontinus Type A, fin length

Measurements (nm) of larvae of Pontinus Type B. (Specimens between dashed lines
are undergoing notochord flexion. )

Standard
length

Snout-
distance

Head
length

Snout
length

Eye
diameter

Gody
depth

Pectoral
fin
length

Pectoral
fin base
depth

Pelvic

fin

length

Snout-anal

fin
distance

5.0

3.0

2.2

0.70

0.6S

2.3

1.7

0.85

0.96

3.0

5.5

3.8

2.5

0.73

0.70

2.5

1.8

1.0

1.0

3.S

6.0

4.0

2.8

0.82

0.78

2.8

1.8

..........

,......-

-........_

8.7

5.9

3.8

0.98

1.2

3.5

2.1

1.2

1.7

6.0

'10.5

7.0

4.8

1.3

1.5

4.2

2.9

1.6

2.2

7.4

•11.2

7.7

4.7

1.1

1.6

4.7

2.7

1.7

2.1

8.2

•12.7

8.5

5,7

1.4

1.8

5.2

3.5

1.8

2.7

9.1

•13.7

9.5

5.8

1.4

1.9

5.8

3.9

1.9

2.8

10.0

'14.0

9.5

6.2

1.7

2.2

5,4

4.0

1.8

3,0

10.0

•15.4

10.0

6.2

1.5

2.2

6.2

4.4

2.1

3.3

10.6

•17.1

11.3

6.7

1.5

2.5

7.2

4.6

2.3

3.8

11,8

•22.0

13.6

8.8

1.9

2.9

8.8

6.2

2.9

4.3

14,7

'23.1

14.9

9.6

2.2

3.3

9.0

6.2

2.9

4.8

15.9

•Pelagic Juvenile,

64

B

Figure 37.— Developmental series of Pontinus Type B. A. 5.0-mm larva; B. 8.7-mm larva; C. 15.4-imn pelagic juvenile.

65

averages 26% and 32%, respectively. The pectoral fin
base is slightly deeper in Pontinus Type B compared
with Pontinus Type A and like the latter becomes rela-
tively shallower as development proceeds. The fin base
averages 17% of the body length in Pontinus Type B lar-
vae undergoing notochord flexion, 16% in postflexion lar-
vae, and 14% in pelagic juveniles. In comparable stages
of Pontinus Type A, the averages are 15%, 14%, and 13%.
The shape of the fin in Pontinus Type B is aliform as it is
in Pontinus Type A. The sequence of ossification of fin
elements and rays could not be determined because of
calcium leaching in the small larvae during preservation
and storage. In six larger larvae and pelagic juveniles
which were stained, all had 18 pectoral rays. The dorsal,
anal, and pelvic fins were well formed in the smallest lar-
vae available. The counts of the stained specimens were
D. XII, 9; A. Ill, 5; P. I, 5. Three superior and two in-
ferior hypural elements develop in the caudal fin; there
are 8 -(- 7 principal and 5-7 -I- 5-7 procurrent rays. All
stained larvae had 24 vertebrae. Gill rakers of the stained
pelagic juveniles ranged from 5 to 6 on the upper limb
and 9 to 11 on the lower limb of the first arch. There were
7 branchiostegal rays.

The absence of specimens of Pontinus Type B smaller
than 5.0 mm precludes knowledge of the pigment pat-
tern of early larval stages. Pigmentation of larvae larger
than this and pelagic juveniles is similar to that in lar-
vae and pelagic juveniles of Pontinus Type A, but there
are distinctive differences that allow separation of the
two forms.

Several melanophores are present at the ventral mid-
line posterior to the developing anal fin. These remnants
of the ventral midline series are not found in larvae larger
than 8.7 mm. Another component of the original pig-
ment pattern, the melanistic shield covering the gas
bladder, is already masked by trunk musculature in the
5.0-mm larva and becomes further embedded with con-
tinued development.

The pattern on the pectoral fins is markedly different
from that on the fins of Pontinus Type A larvae. On the
5.0-mm specimen a pigment streak extends obliquely
ventrad from the dorsal tip of the fin blade to the mid-
region of the fin. Also, a small blotch is present along the
base of the six most ventral rays and one is present on the
medial surface of the fin base in this region. The medial
blotch is not present in larvae larger than 6.0 mm;
however, the ventral blotch at the base of the fin blade
extends dorsad with further development and ul-
timately connects with the dorsal streak to produce a
complete bar in larvae longer than 10.0 mm. This ob-
lique bar remains a prominent feature throughout the
larval and pelagic juvenile stages. Other fin pigment pres-
ent in the 5.0-mm larva is a patch of melanophores at
the base of each pelvic fin. It remains in larger larvae and
in pelagic juveniles.

Pigment is also present in 5-mm larvae on the spinous
dorsal fin. The blotch on the spinous dorsal membrane
extends from the 3rd or 4th spine to the 6th, 7th, or 8th
spine and remains throughout later developmental
stages. It enlarges slightly with development and covers

the membrane from the Ist to 10th rays in the 23.1-mm
pelagic juvenile. A portion of this blotch from the 6th to
10th ray is darker than the background and stands out as
a black spot. This same pattern is present on the spinous
dorsal fin of a 39.0-mm benthic juvenile of P. sierra.

The dorsal pigment saddles appear much earlier than
in larvae of Pontinus Type A and are distinctly different
in form. In 5-mm larvae of Pontinus Type B, small
blotches are present on either side of the dorsal midline
at the anterior end of the spinous dorsal fin and at the
posterior region of the fin. In 8-mm larvae the anterior
saddle has enlarged to occupy the region along the an-
terior one-third of the spinous dorsal and has enlarged
ventrad to the head and forward over the nape. A narrow
space separates the anterior saddle from the posterior
saddle which now extends along the posterior two-thirds
of the spinous dorsal and ventrad about halfway to the
lateral line. A third pigment saddle is beginning to form
on either side of the soft dorsal fin. In 10-mm larvae the
posterior saddle of the spinous dorsal fin extends to the
lateral line and the one at the soft dorsal extends to the
horizontal septum. In 14-mm larvae the latter saddle ex-
tends ventrad to the anal fin where it meets a blotch of
pigment on the membrane of the soft anal rays. Anterior
to this bar three distinctive blotches lie along the
horizontal septum. At about 17 mm a blotch forms on
each side of the caudal peduncle, and in the largest
pelagic juvenile (23.1 mm) it has connected with dorsal
midline pigment to form a saddle. Also, in this largest
specimen a blotch is present over each side of the hy-
pural region. The pattern of pigment saddles and
blotches in the largest pelagic juveniles matches closely
the pattern found in a 39.0-mm benthic juvenile of P.
sierra.

Head pigment first appears in 8-mm larvae on the dor-
sal and lateral surfaces of the brain and remains
throughout the larval period. In 10-mm larvae, pigment
appears along the basal region of the branchiostegal rays
and on the posterior region of the opercle. Also, a blotch
of pigment is present on the cheek ventroposterior to the
eye. In the largest pelagic juvenile additional pigment
appears anterior to the eye.

Distribution. — Pontinus sierra is known to range from
the Gulf of California to Peru. Pelagic juveniles of Ponti-
nus Type B were taken on two midwater trawl stations in
the southern Gulf of California and on a single midwater
trawl station on the outer coast between Magdalena Bay
and Cape San Lucas (Fig. 38). A wide latitudinal gap
separates these occurrences from the remainder of the
pelagic juveniles and larvae which were taken off the
Central and South American coasts, as far south as the
equator. They appear to be more restricted to coastal
waters than are larvae and pelagic juveniles of Pontinus
Type A and there were no occurrences at the Galapagos
Islands.

Larvae and pelagic juveniles of Pontinus Type B are
rare compared to catches of Pontinus Type A. Of the 23
total positive hauls, 17 contained a single specimen and 6
contained two specimens.

66

WPI'IM'I'I'

"'■I'i'i'l I, I, Li. I.I, I, I, I, I

.i.l'i.i'i

1. 1. I.I. I

Figure 38. — Collections of larvae and pelagic juveniles of Pontinus
Type B from various sources. Open circles, Scripps Tuna
Oceanography cruises, closed circles, EASTROPAC Expedition;
open squares, Scripps Institution of Oceanography iish collection.

Ectreposebastes Garman

Literature. — In their review of the scorpaenid sub-
family Setarchinae, Eschmeyer and Collette (1966) gave
a brief description of two larval specimens (15 and 16 mm
SL) of E. imus. Illustrations of the larvae have not ap-
peared in the literature.

Distinguishing features. — Larvae of Ectreposebastes
can be distinguished from those of other eastern Pacific
scorpaenid genera by a combination of characters. They
have enormous pectoral fins. The depth of the fin base is
19 to 22% of the body length while in the larvae of other
tropical scorpaenid genera, except Scorpaenodes, it
ranges from 12 to 18% of the body length. In Scor-
paenodes, the depth of the pectoral fin base ranges from
16 to 21% of the body length over most of the larval
period. Larvae of Scorpaenodes have a bifurcate parietal
ridge whereas Ectreposebastes larvae have a spikelike
spine at the terminus of each parietal ridge. Also, the
pectoral fins of Ectreposebastes are longer than in other
scorpaenid genera. In larvae which have completed
notochord flexion, the pectorals reach or extend beyond
the base of the caudal fin. Also, larvae of Ectreposebastes
are deeper-bodied than other scorpaenid genera in the
eastern Pacific. The rays of the dorsal and anal fins are
highly elongate and the pterygiophores and overlying
muscles appear to protrude from the body profile. Esch-
meyer and Collette (1966) considered Ectreposebastes to
be a monotypic genus of the tropical Atlantic and Pacific
but, in a recent paper, Eschmeyer and Randall (1975)
suggested that E. niger (Fourmanoir) may be a distinct
species. Only a single larval form was recognizable and is
herein referred to as E. imus.

Ectreposebastes imus Garman, Figure 39

Literature. — As above.

Distinguishing features. — Size at hatching is un-
known; the smallest larva in our collection is 2.8 mm.
The larvae attain a larger size than any other known
species in the family. A 28.2-mm specimen is just begin-
ning to develop juvenile pigmentation on the head.

Larvae of E. imus are the deepest- and narrowest-
bodied of all known eastern Pacific scorpaenid larvae.
Body depth averages 31% of body length in the preflex-
ion larvae and increases gradually thereafter to 55% in
the 28.2-mm transforming specimen (Table 38). The gut
has a compact anterior mass with a long intestinal sec-
tion extending posteriad to the anus. Snout-anus dis-
tance is large; it averages 53% of the body length in
preflexion larvae and increases thereafter to a maximum
of 76% in the 23.0-mm specimen.

The head is large and develops a pair of spikelike
parietal spines, each with a short posteriorly directed
nuchal spine behind it. Head length averages 34% of the
body length before and during notochord flexion, and
averages 38% at completion of flexion. The eyes are small
and undergo a relative diminution during the larval
period. Eye diameter averages 32%i of head length before
and during notochord flexion and is gradually reduced to
a minimum of 22% in the 23.0-mm larva. Snout length
averages 33% of head length over the entire larval period.

Larvae of E. imus have distinctive fins. The pectoral
fins are the longest of any known scorpaenid. Pectoral fin
length averages 33% of the body length in preflexion lar-
vae, 38%i during flexion, and increases to a maximum of
57% of the body length in the transforming specimen.
They extend to or beyond the hypural elements in larvae
6.7 mm and longer. Also, the fin base is unusually deep;
it ranges from 19 to 22% of the body length during the
larval period. Although there were too few larvae in the
series to permit staining, the development of fin rays can
be observed on unstained specimens. The full com-
plement of 18 to 20 pectoral rays is present in the
smallest larva (3.8 mm) in our collection. The pelvic fin
is moderate in size and is first apparent in the 5.5-mm
larva where its length is 9%. of the body length. Fin length
increases gradually during the remaining larval period to

Table 38. Measurements (rm) of larvae of Ectreposebastes imus. [Specirrens between dashed
lines are undergoing notochord flexion.)

Standard
length

Snout-
distance

Head
length

Snout
length

Eye

diameter

Body
depth

Pectoral
fin

length

Pectoral
fin base

depth

Pelvic

fin

length

Snout-anal

fin

distance

2.8

1-6

1.0

0.31

0.31

1.0

0.S8

0.54

.

.

3.2

1.8

1.1

0-38

0.38

1.1

1.2

0.68

-

3.4

1.9

1.2

0.43

0.42

1.4

1.3

0.76

3.8

1.9

1.2

0.43

0.38

1.4

1.2

0.72

*-^ J

2.0

1.4 J

0.45

0.40

1.5

1.3

0.90
.........

0.50

2.2

"'"i'i"'

6.5

3.2

"'i'i'"

1.9

"'i'.i'

0.65

"o'.Te"'

0.58

"""o'-Vf"'

2.2

"'i'.i'

2.1

2.3

1.3

0.66

3.4

6.7

4.2

2.7

0.88

0.80

2.8

3.3

1.5

1.2

4.3

9.8

6-5

3.8

1.2

1.1

4.6

4.2

2.1

2.2

6.7

11.3

7.6

4.6

\.z

1.2

5.4

5.0

2.5

2.5

7.7

23.0

17.6

9.2

2.7

2.0

11.2

n.3

4.4

5.0

17.7

•28.2

19.9

10.1

3.3

2.5

16.6

16.0

5.8

7.9

20.3

•Transforming specimen.

67

Figure 39. — Developmental stages of Ectrepoeebastea imus. A. 6.7-inin larva; B. 23.0-inm larva; C. 28.2-mm transform-
ing specimen £rom stomach of Alepi»aurus (note teeth lacerations).

68

a maximum of 28% of the body length in the trans-
forming specimen. The full complement of I, 5 pelvic
rays is present in the 6.7-mm larva.

The dorsal and anal fins are beginning to develop in
the 4.'2-mm larva and the full complements of XII, 10(9)
dorsal rays and III, 6(5) anal rays are present in the 6.7-
mm larva. In the 23.0-mm larva the third dorsal spine is
2.5% of the body length.

The hypural anlagen and the principal caudal rays
have begun to develop in the 3.8-mm larva. The full com-
plements of 3 -(- 2 hypurals and 8 -(- 7 caudal rays are visi-
ble in the 5.5-mm larva. The procurrent rays are begin-
ning to form at 5.5 mm, however, the full complement of
5-6 + 6-7 rays is not present until late in the larval
period.

In the smallest larvae, melanistic pigment is present
on the dorsal surface of the developing gas bladder, along
the ventral surface of the gut, and on the pectoral fins.
The series of melanophores along the ventral midline of
the tail contains 11 to 14 pigment spots which disappear
before the beginning of notochord flexion. The ventral
gut pigment is lost before the larvae reach 4.0 mm and
that above the gas bladder becomes obscured by trunk
musculature. In the smallest larvae, the pectoral fin
membrane is covered solidly with small melanophores.
When the larvae reach 5.5 mm, there is an unpigmented
region at the base of each fin and, with continued develop-
ment, the melanistic zone becomes more distally located.
In the 23.0-mm larva, the distal half of the fin is pigment-
ed and in the transforming specimen the distal third of
each fin is pigmented. In the latter specimen, the fine
melanophores that will eventually cover the entire body
are beginning to appear on the head and upper trunk.

Distribution. — According to Eschmeyer and Collette
(1966) and Eschmeyer and Randall (1975), E. imus is
known from the eastern and western Atlantic, the
southeastern Pacific off the Galapagos Islands, and Peru,
Hawaii, and Japan. Specimens in this study were taken
from about lat. 2° to 6°N off the coast of Colombia and
Ecuador, at the equator between long. 100° and 127°W,
and at the Hawaiian Islands (Fig. 40). Larvae of £. imus
are extremely rare in plankton collections. Of the 15
specimens available to us, 8 came from EASTROPAC 1-
m plankton hauls, 6 were taken by midwater trawls, and
the transitional specimen came from an Alepisaurus
stomach.

ACKNOWLEDGMENTS

Figure 40. — Collections of larvae of Ectreposebastes imiu from
EASTROPAC expeditions, Scripps Tuna Oceanopaphy cruises,
Scripps Institution of Oceanography fish collection, and the Univer-
sity of Hawaii.

ferring specimens to our laboratory. We thank John
Fitch (California Department of Fish and Game, Long
Beach) and Carl L. Hubbs and Richard Rosenblatt (SIO)
for sharing with us their extensive knowledge of scor-
paenid fishes. We are especially indebted to Lo-chai
Chen (San Diego State University), William Eschmeyer
(California Academy of Sciences), and Jurgen Westr-
heim (Fisheries Research Board of Canada, Nanaimo)
for reviewing the manuscript and offering valuable sug-
gestions for improving it. Eschmeyer's comments were
particularly extensive on Trachyscorpia and North At-
lantic Sebastes and were essential to the writing of these
sections. From the Southwest Fisheries Center, NMFS,
La Jolla, Amelia Gomes and Betsy Stevens provided
technical help during the course of the study; George
Mattson prepared the illustrations in Figures la-c, 2c, 3,
4b, 5, 7, and 24 through 29; and Ken Raymond and Henry
Orr drafted Figures 18, 19, 30, 36, and 38. Special thanks
goes to John LaGrange and Richard Pleasant for supply-
ing live rockfish larvae and to David Kramer (NMFS,
LaJolla) for aiding in the culturing of these larvae. Pam-
ela Moser generously gave her time in typing parts of the
manuscript.

This paper is dedicated to the memory of Don Dockins
who, as a worker in the fish collection of Scripps In-
stitution, untiring colleague on countless collecting trips,
and friend of the senior author, aided greatly in the com-
pletion of the paper.

This study was facilitated by the generous efforts of
many people. For the loan of valuable specimens we are
indebted to Robert Lavenberg, Los Angeles County
Museum; Richard Rosenblatt, SIO; Erik Bertelsen,
University Museum, Copenhagen; George Kelley,
Thomas McKenney, and Ruth Stoddard, Northeast
Fisheries Center, NMFS; Thomas Clark, University of
Hawaii; and C. R. Hitz, Northwest Fisheries Center,
NMFS, Seattle. Jerry Newmann, LACM, and Joseph
Copp, SIO, provided valuable help in locating and trans-

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70

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71

388. Proceedings of the first U.S. Japan meeting on aquaculture at
Tokyo, Japan. October 1819, 1971. William N. Shaw (editor). (18
papers, 14 authors.) February 1974, iii + 133 p. For sale by the
Superintendent of Documents, U.S. Government Printing Office,
Washington, D.C. 20402.

389. Marine flora and fauna of the northeastern United States.
Crustacea: Decapoda. By Austin B. Williams. April 1974, iii + ."JO p.. Ill
figs. For sale by the Superintendent of Documents, U.S. Government
Printing Office,' Washington, D.C. 20402.

390. Fishery publications, calendar year 1973: Lists and indexes. By
Mary Ellen Engett and Lee C. Thorson. September 1974, iv + 14 p., 1 fig.
For sale by the Superintendent of Documents, U.S. Government Printing
Office, Washington, DC. 20402.

391. Caianoid copepods of the genera Spinocalanus and Mimncalanua
from the central Arctic Ocean, with a review of the Spinocalanidae. By
David M. Damkae'. June 1975, x + 88 p., 225 figs, 4 tables. For sale

by the Superintendent of Documents. U.S. Government Printing Office,
Washington, DC. 20402.

392. Fishery publications, calendar year 1974: Lists and indexes. By
Lee C. Thorson and Mary Ellen Engett. June 1975, iv + 27 p.. 1 fig.

393. Cooperative Gulf of Mexico estuarine inventory and study— Texas:
Area description. By Richard A. Diener. September 1975, vi + 129 p.,
.55 figs., 26 tables,

.394 Marine Flora and Fauna of the Northeastern United States. Tar-
Hinrada Bv l.cland W. Pollock. Mav 1976, iii + 25 p., figs. For sale
l)v ihc .Superintendent ol Doiumcnls. U.S. Government Printing Office.
Wii.shingliin. D.C. 20402.

.395. Report of a colloquium on larval fish mortality studies and their
relation to fishery research, January 1975. By John R. Hunter. May
1976, iii + 5 p. For sale by the Superintendent of Documents, U.S.
Government Printing Office, Washington, DC. 20402.

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