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NOAA Technical Report
cular 430
Guide to Identification of
Some Sculpin (Cottidae)
Larvae from Marine and
Brackish Waters off Oregon
and Adjacent Areas in
The Northeast Pacific
Sally L. Richardson and Betsy B. Washington
January 1980
U.S. DEPARTMENT OF COMMERCE
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NOAA Technical Report NMFS Circular 430
Al MOSp,
NT Of C
Guide to Identification of
Some Sculpin (Cottidae)
Larvae from Marine and
Brackish Waters off Oregon
and Adjacent Areas in
the Northeast Pacific
Sally L. Richardson and Betsy B. Washington
January 1980
2?
(X
o
U.S. DEPARTMENT OF COMMERCE
Juanita M. Kreps, Secretary
National Oceanic and Atmospheric Administration
Richard A. Frank, Administrator
National Marine Fisheries Service
Terry L. Leitzell, Assistant Administrator for Fisheries
For Sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, DC. 20402 - Stock No. 003-017-00472-4
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publication.
CONTENTS
Introduction 1
Methods 2
Useful characters 3
Order of presentation • 4
Artedius harringtoni (Starks) 4
Artedius Type 2 8
Clinocottus acuticeps (Gilbert) 12
Cottidae Type 1 15
Cottidae Type 2 IV
Cottidae Type 3 18
Trigolops sp 20
Chitonotus pugetensis (Steindachner) 22
Cottus asper Richardson 24
Enophrys bison (Girard) 25
Icelinus spp 27
Leptocottus armatus Girard 31
Paricelinus hopliticus Eigenmann and Eigenmann 32
Hemilepidotus hemilepidotus (Tilesius) 34
Hemilepidotus spinosus (Ayres) 37
Nautichthys oculofasciatus (Girard) 40
Radulinus asprellus Gilbert 42
Radulinus boleoides Gilbert 44
Rhamphocottus richardsoni Giinther 46
Scorpaenichthys marmoratus (Ayres) 48
Cottoid Type A 51
Notes on other northeast Pacific cottid larvae 53
Acknowledgments 54
Literature cited 55
Figures
1. Larvae of Artedius harringtoni 6
2. Larvae of Artedius harringtoni 7
3. Larvae of Artedius Type 2. The second figure from the top is a dorsal view showing gut diverticula ... 9
4. Larvae of Artedius Type 2 10
5. Larvae of Clinocottus acuticeps. The second figure from the top is a ventral view showing hindgut
diverticula 12
6. Young of Clinocottus acuticeps 13
7. Larvae of Cottidae Type 1 16
8. Larvae of Cottidae Type 2 17
9. Larvae of Cottidae Type 3 19
10. Larvae of Triglops sp 21
11. Larvae of Chitonotus pugetensis 22
12. Young of Chitonotus pugetensis 23
13. Larvae of Cottus asper 25
14. Young of Enophrys bison 26
15. Larvae of Icelinus spp 28
16. Young of Icelinus spp 29
17. Young of Icelinus spp 30
18. Larvae of Leptocottus armatus 31
19. Larvae of Paricelinus hopliticus 33
20. Young of Paricelinus hopliticus 34
21. Larvae of Hemilepidotus hemilepidotus 35
22. Young of Hemilepidotus hemilepidotus 36
23. Larvae of Hemilepidotus spinosus 38
24. Young of Hemilepidotus spinosus 39
25. Larvae of Nautichthys oculofasciatus 41
26. Larvae of Radulinus asprellus 43
27. Young of Radulinus asprellus 44
iii
28. Larva of Radulinus boleoides 45
29. Young of Rhamphocottus richardsoni 47
30. Larvae of Scorpaenichthys marmoratus 48
31. Young of Scorpaenichthys marmoratus 49
32. Larvae of Cottoid Type A 52
Tables
1. Species of adult cottids whose range of distribution reportedly includes marine or brackish waters off
Oregon and their meristics 2
2. Groupings of cottid larvae described in this guide based on preopercular spines and pigment pat-
terns 5
3. Meristics from larvae of Artedius harringtoni 5
4. Measurements (mm) of larvae of Artedius harringtoni 5
5. Meristics from larvae of Artedius Type 2 8
6. Measurements (mm) of larvae of Artedius Type 2 8
7. Meristics from young of Clinocottus acuticeps 14
8. Measurements (mm) of larvae of Clinocottus acuticeps 14
9. Meristics of larvae of Cottidae Type 1 15
10. Measurements (mm) of larvae of Cottidae Type 1 , 15
11. Meristics from larvae of Cottidae Type 2 18
12. Measurements (mm) of larvae of Cottidae Type 2 18
13. Meristics from larvae of Cottidae Type 3 20
14. Measurements (mm) of larvae of Cottidae Type 3 20
15. Meristics from larvae of Triglops sp 20
16. Measurements (mm) of larvae of Triglops sp 20
17. Meristics from larvae of Chitonotus pugetensis 23
18. Measurements (mm) of larvae of Chitonotus pugetensis 24
19. Meristics from larvae of Cottus asper 24
20. Measurements (mm) of larvae of Cottus asper 24
21. Meristics from larvae of Enophrys bison 25
22. Measurements (mm) of larvae of Enophrys bison 27
23. Meristics from larvae of Icelinus spp 30
24. Measurements (mm) of larvae of Icelinus spp 31
25. Meristics from larvae of Leptocottus armatus 32
26. Measurements (mm) of larvae of Leptocottus armatus 32
27. Meristics from young of Paricelinus hopliticus 32
28. Measurements (mm) of young of Paricelinus hopliticus 33
29. Meristics from young of Hemilepidotus hemilepidotus 34
30. Measurements (mm) of young Hemilepidotus hemilepidotus 36
31. Meristics from young of Hemilepidotus spinosus 40
32. Measurements (mm) of young of Hemilepidotus spinosus 41
33. Meristics from larvae of Nautichthys oculofasciatus 42
34. Measurements (mm) of larvae of Nautichthys oculofasciatus 42
35. Meristics from young of Radulinus asprellus 42
36. Measurements (mm) of young of Radulinus asprellus 45
37. Meristics from larvae of Radulinus boleoides 45
38. Measurements (mm) of larvae of Radulinus boleoides 45
39. Meristics from larvae of Rhamphocottus richardsoni 46
40. Measurements (mm) of larvae of Rhamphocottus richardoni 46
41. Meristics from young of Scorpaenichthys marmoratus 50
42. Measurements (mm) of young of Scorpaenichthys marmoratus 50
43. Meristics from larvae of Cottoid Type A 51
44. Measurements (mm) of larvae of Cottoid Type A 52
45. Additional northeast Pacific cottid species for which larvae are known 53
IV
Guide to Identification of Some Sculpin (Cottidae)
Larvae from Marine and Brackish Waters off
Oregon and Adjacent Areas in the Northeast Pacific
Sally L. Richardson1 and Betsy B. Washington2
ABSTRACT
Knowledge of identification of cottid larvae taken in plankton collections off Oregon is sum-
marized. Included are descriptions of 21 kinds of cottid larvae representing at least 14 of the 40 genera
of cottids reported to occur in the northeast Pacific between Baja California and the Aleutian Islands.
Larvae of Artedius harringtoni, Chitonotus pugetensis, Clinocottus acuticeps, Icelinus spp.,
Paricelinus hopliticus, Radulinus asprellus, R. boleoides, and four unidentified types are described
for the first time. A larval series of a second kind of Artedius is described with a discusion of problems
of specific identification. New and comparative information is provided for larvae of Cottus asper,
Enophrys bison, Hemilepidotus hemilepidotus, H. spinosus, Leptocottus armatus, Nautiehthys oeulo-
fasciatus, Rhamphocottus richardsoni, Scorpaenichthys marmoratus, and Triglops sp.
Each descriptive account contains illustrations, a literature summary, distinguishing features
(including size at hatching and transformation, pigmentation, head spination, general morphology),
and a discussion of identification problems. Characters found to be most useful in distinguishing cot-
tid larvae include morphology, pigmentation, head spination, and meristics.
Literature summaries are also provided for northeast Pacific cottid larvae in 8 additional genera:
Blepsias, Dasycottus, Gilbertidia, ?Malacoeottus, Myoxocephalus, Oligocottus, Orthonopias,
Psychrolutes.
INTRODUCTION
The cottids are one of the most speciose and diverse
groups of fishes inhabiting coastal waters of the north-
east Pacific Ocean with 90 species in 40 genera reported
to occur from Baja California to the Aleutian Islands
(Howe and Richardson1). The range of 36 of these
species in 20 genera reportedly includes waters off
Oregon (Table 1). Adult cottids range in size from the 5
cm Artedius meanyi, Clinocottus acuticeps, and
Oligocottus rimensis to the 76 cm Myoxocephalus poly-
acanthoceihalus and Scorpaenichthys marmoratus
(Hart 1973). Larger forms in the genera Gymno-
canthus, Hemilepidotus, Hemitripterus, Melletes, and
Myoxocephalus are fished commercially by the
U.S.S.R. and Japan (Gorbonova 1964; Fedorov 1973).
Off Oregon several species are taken frequently by
sports fishermen, e.g., Enophrys bison, Hemilepidotus
hemilepidotus, H. spinosus, Leptocottus armatus, Scor-
paenichthys marmoratus (Beardsley and Bond 1970).
'School of Oceanography, Oregon State University, Corvallis, Oreg.;
present address: Gulf Coast Research Laboratory, East Beach Drive,
Ocean Springs, MS 39564.
'School of Oceanography, Oregon State University, Corvallis, OR
97330.
Howe, K., and S. L. Richardson. 1978. Taxonomic review and
meristic variation in marine sculpins (Osteichthys: Cottidae) of the
northeast Pacific Ocean. Final Report, NOAA NMFS Contract No. 03-
78-M02-120, 1 January 1978 to 30 September 1978, 142 p. Northwest
and Alaska Fisheries Center, National Marine Fisheries Service, NOAA,
2725 Montlake Blvd. East, Seattle, WA 98102.
Most, if not all cottid larvae hatch from demersal eggs
~(about)l-4 mm in diameter (Breder and Rosen 1966)
and range from approximately 3 to 12 mm at hatching.
They are usually pelagic and some are primarily
neustonic, i.e., occurring at the surface of the water
column, such as Scorpaenichthys marmoratus and
Hemilepidotus spinosus (unpubl. data). Most cottid lar-
vae probably transform to juveniles by ~20 mm SL.
Cottid larvae are collected frequently in the coastal
zone of temperate and boreal areas of the northeast
Pacific but they are usually not abundant. In a l'/a yr
survey off Oregon they occurred in 80' 7c of the samples
but comprised only 8ch of the fish larvae captured in a
coastal assemblage of larval fishes 2-28 km offshore
(Richardson 19774; Richardson and Pearcy 1977). They
were taken in all months of the year in that study.
A wide variety of larval forms is demonstrated within
the cottid family. Identification of the larvae is often dif-
ficult because of the large number of species in the fami-
ly, because of difficulty in establishing developmental
series from the small numbers of larvae in plankton
samples, and because relatively few species have been
described in detail.
This report summarizes knowledge of identification of
cottid larvae taken primarily in plankton and neuston
^Richardson, S. L. 1977. Larval fishes in ocean waters off Yaquina
Bay, Oregon: abundance, distribution, and seasonality January 1971 to
August 1972. Oreg. State Univ., Sea Grant Coll. Prog. Publ. No.
ORESU-T-77-003, 73 p.
Table 1. — Species of adult cottids whose range of distribution reportedly includes
marine or brackish waters off Oregon and their meristics.1 Branchiostegal rays num-
ber 6 unless otherwise noted.
Dorsal
Dorsal
fin
Anal
fin
soft
fin
Pectoral
Pelvic
Total
Species
spines2
rays2
rays-
fin rays
fin rays
vertebrae
Artedius eorallinus
VIII-IX
15-16
12-13
14-16
3
31-33
Artedius fenestralis
VIII -IX
16-18
12-14
14-16
3
32-35
Artedius harringtoni'
IX
15-18
10-14
13-15
3
32-34
Artedius lateralis
VII-X
15-17
12-14
14-16
3
32-34
Artedius meanyi
IX-X
14-17
10-12
14-16
2-3*
33-35
Artedius notospilotus
IX-X
14-16
11-13
14-17
3
32-34
Ascelichthys rhodorus
VII-X
17-19
13-16
16-18
0
33-36
Blepsias cirrhosus
VI-VIII
20-24
18-21
11-13
3
37-38
Chitonotus pugetensis
VIII- XI
14-17
14-17
16-18
2-3*
35-36
Clinocottus acuticeps
VII-IX
13-17
9-13
13-15
3
31-33
Clinocottus embryum
vni-x
14-17
9-12
12-15
3
33-34
Clinocottus globiceps
vm-x
13-17
11-12
13-15
3
32-34
Clinocottus recaluus
VIII-IX
14-16
9-13
13-15
3
32-33
Cottus aleuticus
VIII-X
16-20
12-16
13-16
4
34-39
Cottus asper"
VII-XI
18-21
14-18
14-17
4
34-39
Enophrys bison
VII-IX
9-13
8-10
15-17
3-4*
29-31
Hemilepidotus hemilepidotus
X-XIII
18-20
13-16
15-17
4
35-37
Hemilepidotus spinosus
XXI
18-20
14-16
14-16
4
35-37
Icelinus burchami
XXI
16-18
12-14
16-19
2
35-37
Icelinus filamentosus
X-XII
15-17
13-16
16-18
2
34-37
Icelinus oculatus
XI
15-17
13-14
17
2
37
Icelinus tenuis
IX-XI
16-18
14-17
15-17
2
37-39
Jordania zonope
XVII-XVIII
15-18
22-24
13-15
4-5*
46-48
Leptocottus armatus
VI-VIII
15-20
15-20
17-20
4
35-39
Nautichthys oculofasciatus
VIII-IX
27-30
16-21
13-14
3
40-41
Oligocottus maculosus
VIII-IX
15-18
12-14
12-15
3
33-34
Oligocottus rimensis
VIII-X
16-19
13-15
13-15
3
34-37
Oligocottus snyderi
VII-IX
17-20
12-15
12-15
3
34-37
Paricelinus hopliticus
xii-xm
19-20
23-24
14-15
5
42
Psychrolutes phrictus4
VII-IX
19-20
12-14
22-26
3
33-35
Radulinus asprellus
VIII- X
20-23
21-25
17-20
3
38-39
Radulinus boleoides
VIII-XI
20-22
21-23
18-20
3
39-40
Rhamphocottus richardsoni
VII-IX
12-14
6-8
14-16
3*-4
26-28
Scorpaenichthys marmoratus
VIII- XII
15-19
11-14
14-16
4-5*
35-37
Synchirus gilli
VIII-X
19-21
18-21
21-24
3
38-39
Zesticelus profundorum
Y VII
10-13
8-11
19-21
2*-3
25-26
'From compilation by Howe and Richardson 1978 (see text footnote 3) which incorporated
data from literature sources and original counts.
"Includes all fin ray elements whether or not they arise from the same pterigiophore.
'Includes hypural.
'Has 7 branchiostegal rays instead of the usual 6.
'May have 6 or 7 branchiostegal rays.
*Most frequent count.
collections off the Oregon coast. Included are descrip-
tions and illustrations of complete or partial
developmental series of 21 kinds of cottid larvae. These
represent all but one form of cottid larvae from Oregon
waters for which identified (as species or type) material
is available in the Oregon State University Larval Fish
Reference Collection. Although keys are not provided,
larvae are grouped together on the basis of common lar-
val characters. Occurrences, including seasonality, in-
shore-offshore distribution and relative abundance of
most of these cottid larvae off Oregon have been
reported by Richardson (see footnote 4) and Richardson
and Pearcy 1977. However, relevant new information on
occurrences is presented where appropriate. Knowledge
of other cottid larvae from the northeast Pacific is sum-
marized from the literature and personal observation.
This paper provides a basis for comparison with ad-
ditional cottid species whose larvae remain to be iden-
tified and described. It is primarily descriptive in
nature. Examination of relationships withinin the
northeast Pacific Cottidae based on larval morphology
is presented elsewhere (Richardson in press).
METHODS
Specimens came primarily from over 1,000 plankton
and neuston samples collected off Oregon since 1969,
which contained over 2,000 cottid larvae. During the
course of working out identifications additional mate-
rial was examined from plankton collections taken off
northern California, Washington, Puget Sound, British
Columbia, and Alaska (see Acknowledgments). Some
transforming and juvenile specimens collected from
Oregon tidepools were also examined. All material had
been preserved in either 5 or 10% Formalin and some
specimens had been transferred to 36 or 40% isopropyl
alcohol.
Developmental terminology generally follows
Ahlstrom et al. (1976) where the larval period is
separated into three stages, preflexion, flexion, and
pcstflexion, on the basis of the relative position of the
notochord tip. Some subjectivity is involved in deciding
the exact size at which the notochord begins to flex and
when it is in a fully flexed position, but we tried to be
consistent in judgment in all of the series. The transi-
tion from postflexion larva to juvenile could not be
precisely defined in most series because of lack of
specimens in the critical size range. Generally, the
transformation to juvenile is accompanied by an in-
crease in body pigmentation, scale formation, and com-
pletion of development of all fin rays. We refer to fish as
postflexion larvae until they have attained an adult
body form which characterizes them as juveniles.
Counts of various body parts were usually made on
unstained specimens. When adequate material was
available, a developmental series or several specimens
were stained using the method of Taylor (1967). In un-
stained specimens fin rays were counted if any elements
were visible. In stained specimens, fin rays were counted
if any part absorbed stain. Fin ray counts include all ray
elements whether or not they arise from the same
pterygiophore. Vertebral counts include the urostyle.
Measurements were made following definitions given
by Richardson and Laroche (1979) except: Body length
= snout tip to notochord tip preceding development of
caudal fin [notochord length (NL)], then to posterior
margin of hypural plate [standard length (SL)]. This
applies to all lengths given in this paper unless noted
otherwise. Snout length = horizontal distance from tip
of snout to anterior margin of pigmented region of left
eye. Eye diameter = greatest diameter of pigmented
portion of eye. Preopercular spine length (second spine)
= from tip to basal insertion of core of spine. Head
length is abbreviated as HL.
Head spine terminology in cottid larvae has not been
well defined. Many spines present during larval
development are not present or obvious in adults and
names used for spines in adults are often inconsistent. A
standard terminology for spines in larval cottids, based
on studies of developmental osteology of many species,
remains to be proposed. In this paper, names used have
been generally adapted from those that Richardson and
Laroche (1979) diagramed for larvae and juveniles of
Sebastes spp. for which similar problems exist. Some
deviations from their terminology for Sebastes spp. exist
in cottids. Preopercular spines along the posterior
margin frequently number 4 but sometime may be mul-
tiple (over 5) serrations or small spines, rather than 5 as
in Sebastes spp. Additional terms used here include:
spines in the parietal region (several spines in the
vicinity of the parietal spines of Sebastes spp.,
sometimes called occipital spines in adult cottids);
spines in the posttemporal-supracleithral region
[several spines in the vicinity of the posttemporal and
supracleithral spines of Sebastes spp., the basal origin of
which could not be determined on unstained cottid lar-
vae (not to be confused with spiny scales that develop in
this region along the lateral line)); anterior parietal
spine or bump (apparently on the parietal ridge ante-
rior to the principle parietal spine); midopercular spine
(near the middle of the posterior margin of the opercle).
Illustrations were made with the aid of camera lucida.
They are intended to accurately represent numbers of
myomeres, fin rays including ray elements and bases,
head spines, scales, and ventral midline melanophores.
Identifications in most cases could be made using
adult characters, primarily meristic (Table 1; Howe and
Richardson footnote 3) on the largest larvae in a series
linked together by pigment and head spine patterns.
Additional adult characters used to confirm identifica-
tions included deep emargination between dorsal fin
spines 3 and 4 in Chitonotus pugetensis, attachment of
inner pelvic ray to belly by membrane in Clinocottus
aeutieeps, number of dorsal scale rows and scales above
the lateral line in Hemilepidotus hemilepidotus and H.
spinosus, elongated first dorsal fin in Nautichthys oculo-
fasciatus, and unique shape in Rhamphocottus richard-
soni. Identification of larvae of Radulinus boleoides was
based on their similarity to R. asprellus. Newly hatched,
reared larvae of Hemilepidotus hemilepidotus, Dunn
helped confirm the identification of small larvae in that
series.
The descriptive format is designed to present a con-
cise account of a combination of characters which,
together with meristics (not included in the descrip-
tions), will help to distinguish each of the kinds of lar-
vae. Available literature is summarized for each larval
type and lengths are given for each specimen illustrated.
Distinguishing features in each account include infor-
mation on size at hatching and transformation, pig-
mentation, head spination, and general morphology.
Discussions for each kind of larvae may include taxo-
nomic problems, comparisons with similar species, and
new occurrence data.
USEFUL CHARACTERS
Characters most useful to distinguish cottid larvae
include general shape of the body and snout, unusual
diverticula of the gut cavity, melanistic pigment pat-
terns, head spination, and meristics. Body shape can
range from short and stubby (Artedius harringtoni,
Clinocottus aeutieeps) to moderately long and slender
(Icelinus spp., Triglops sp.). Cottid larvae have a distinc-
tively coiled gut and the hindgut may trail below the
body (Artedius harringtoni, Clinocottus aeutieeps).
Snout to anus length is usually ~ 40-50% SL except in
small (<5 mm) larvae where it may be less. However,
some species have a rather long gut, >60% SL (Clinocot-
'Dunn, J. R. 1973. Unpubl. data. Northwest and Alaska Fisheries
Center, National Marine Fisheries Service, NOAA, 2725 Montlake Blvd.
East, Seattle, WA 98102.
tus acuticeps, Rhamphocottus richardsoni). Small (<6
mm) cottid larvae have a very characteristically pro-
truding snout which, with development, can vary con-
siderably in its degree of pointedness (e.g., compare
Scorpaenichthys marmoratus and Radulinus asprellus).
Several forms have unusual gut diverticula or protru-
sions from the abdominal wall {Artedius Type 2,
Clinocottus acuticeps) which serve as striking
characters.
Pigment patterns can range from darkly pigmented
forms such as Scorpaenichthys marmoratus and
Radulinus asprellus where melanophores cover the entire
body except the tail tip to relatively unpigmented forms
such as Triglops sp. with melanophores only on the head
and gut. In many species (Artedius harringtoni,
Chitonotus pugetensis, Paricelinus hopliticus) the
number, extent, spacing, and shape of the postanal
ventral midline melanophores serve as diagnostic aids,
even though these may decrease in number with develop-
ment. Ranges and frequency of counts of these melano-
phores within developmental periods are useful. The
presence or absence of nape pigment and/or head pig-
ment, particularly in small (<5 mm) larvae, may be
helpful in some cases {Artedius spp., Clinocottus spp.,
Enophrys spp.). The pattern and extent of pigment over
the gut is useful in otherwise lightly pigmented forms. It
may range from relatively heavy pigment over the entire
gut (Paricelinus hopliticus, Triglops sp.) to relatively
light concentrations over the dorsolateral surface only
(Cottidae Type 1, Cottidae Type 2). Sometimes it ap-
pears in distinctive patterns (the bars in Leptocottus ar-
matus, the distinctive round melanophores in Enophrys
bison) although these patterns can be obscured in poorly
or improperly preserved material.
Most cottid larvae develop four spines on the preoper-
cle. However, some forms develop only one prominent
spine (Rhamphocottus richardsoni) while others develop
a large number, up to ~25, of smaller spines (Artedius
spp.) on the preopercular margin. A single parietal and
nuchal spine may develop (Icelinus spp.) or a cluster of
smaller spines (Cottidae Type 1) or no spines at all (Lep-
tocottus armatus). The same situation may prevail in the
posttemporal-supracleithral region. One or more post-
ocular spines may develop (Hemilepidotus spp., Triglops
sp.) or they may be absent (Artedius spp.).
Certain meristic characters (Table 1), particularly
when used in combinations, may be useful in identifying
cottid larvae. While many species have between 32 and
37 vertebrae ( = myomeres), counts at the low or high end
of the range may be helpful, e.g., >40 in Triglops sp. and
Nautichthys oculofasciatus and <31 in Enophrys bison.
The number of pelvic fin rays ranges from 0 to 5 with 3
being the most common count. Relatively few genera
have counts of 2 (e.g., Icelinus) or 5 (e.g., Paricelinus,
Scorpaenichthys) . Pectoral fin ray counts range from 12
to 26 for Oregon species and overlap in many, but counts
at the low or high end of the scale help to single out some
species. Also useful are dorsal and anal fin counts which
for Oregon species range from 5 to 18 dorsal spines, 9 to
30 dorsal rays, and 6 to 25 anal rays. Ranges of these
counts for a particular species are generally 4 or less. The
difference between dorsal (D) and anal (A) soft ray
counts may also be of help which is usually D > A, e.g.,
by 3 to 5 rays in Artedius spp., Clinocottus spp., and
Oligocottus spp. and >9 in Nauthichthys oculofasciatus.
In some species the D and A counts are nearly equal
(Leptocottus) and in some D < A (Paricelinus hopliti-
cus).
Other characters that may be useful include size at
hatching, size at transformation, and cirri patterns. The
size at hatching, e.g., —3 mm in Artedius harringtoni vs.
~12 mm in Hemitripterus americanus (Warfel and Mer-
riman 1944; Okiyama and Sando 1976) and the size at
transformation, e.g., ~9 mm in Enophrys bison and ~16
mm in Chitonotus pugetensis, can serve as dis-
tinguishing characters. Head cirri patterns, which may
be evident in older larvae, may be important in linking
larvae with known juveniles. However, they appear to
develop after the pelagic phase in a number of species,
e.g., Icelinus spp., and may be of limited value.
ORDER OF PRESENTATION
To facilitate use of this guide, the larvae described
herein have been grouped according to certain primary
characters (Table 2). However, a formal dichotomous
key is not provided as many species remain to be de-
scribed and such a key could be misleading. Rather,
groups are formed on the basis of the most outstanding
larval characters. The two major groupings are based on
preopercular spine pattern: group A, numerous (>4);
group B, 4 or less. Within group A, the larvae may be
further distinguished by using a combination of charac-
ters including number of preopercular spines, number of
ventral midline melanophores, number of parietal
spines, and presence or absence of gut diverticulae.
Within group B the larvae may be placed in four sub-
groups on the basis of postanal pigmentation. Within
these four groups, larvae can be further separated using
characters as for group A and relative body shape,
pigmentation, and number of myomeres.
Artedius harringtoni (St arks)
(Figures 1, 2; Tables 3, 4)
Literature. A 4.2 mm SL larva resembling A. harring-
toni was described by Blackburn (1973) as Cottid 6. Lar-
vae of this species were referred to as Artedius sp. 1 by
Richardson (see footnote 4), and Richardson and Pearcy
(1977).
Distinguishing Features. Larvae of A. harringtoni are
probably 3-4 mm long at hatching. The largest pelagic
specimens we collected, ~ 13-14 mm, are beginning to
develop juvenile pigmentation.
A dense concentration of malanophores is present over
the gut. The nape is pigmented in all but a few of the
smallest larvae although this pigment becomes embed-
Table 2. — Groupings of cottid larvae described in this guide based on preopercular spines and pigment patterns.
Ventral1
midline
Preopercular1
melano-
Parietal
Vertebrae2
Additional
spines
phores
spines
(myomeres)
distinguishing characters
A. Preopercular spines numerous ( >4)
Artedius harringtoni
11-21
27-30
0
32-34
Artedius Type 2
13-23
14-24
Multiple
(32-34)
dorsal gut diverticula
Clinocottus acuticeps
8-13
7-9
0
32-33
hindgut diverticula
Cottidae Type 1
19
7
Multiple
(32-35)
Cottidae Type 2
10-11
15-17
1
(32-34)
hindgut bulges
Cottidae Type 3
16-19
4-5
Multiple
(33)
B. Preopercular spines 4 or fewer
1. Postanal pigment absent
Triglops sp.
4
0
1
(47)
dark gut
2. Postanal pigment on ventral midline only
Chitonotus pugetensis
4
41-45
1
35-36
Cottus asper
4
14-17
0
34-39
Enophrys bison
4
11-14
1
29-31
deep bodied
Icelinus spp.
4
10-11
1
33-39
finfold pigment
Leptocottus armatus
4
9-11
0
35-39
pigment bars on gut
Paricelinus hopliticus
4
31-32
1
42
dark gut
3. Postanal pigment primarily on dorsal
and ventral midline on small larvae
Hemilepidotus hemilepidotus
4
14
1
35-37
dorsal pigment interrupted
Hemilepidotus spinosus
4
24-34
1
35-37
dorsal pigment continuous
4. Postanal pigment heavy over body
Nautichthys oculofasciatus
4
N3
B
40-41
large pectoral fins
Radulinus asprellus
4
N
1
38-39
slender bodied
Radulinus boleoides
?
N
?
39-40
Rhamphocottus richardsoni
1
N
1
26-28
deep bodied
Scorpaenichthys marmoratus
4
N
B
35-37
Cottoid Type A
0
N
0
35
globose
'Maximum number at peak of development.
"From Howe & Richardson 1978 (see text footnote 3).
N = not distinguishable; B = bump.
Table 3. — Meristies from larvae of Artedius herringtoni. (Specimen between dashed line is undergoing notochord flexion.)
[* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; N = Not examined.]
Pelvic Ventral Caudal fin rays
Anal Pectoral fin Preoper- midline _ Dorsal Ventral Vertebrae Branchi-
Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal
length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays
3.0 29 N N N N N N N
4.7 12 27 N N N N N N N
6.9 16 30NNNNNNN
7.3 -17-** 13 14 14 16 26 N N N N N N 7
9.3 IX 16 13 14 14 1,3 21 22 N N N N N N 7
11.2* IX 17 14 14 14 1,3 21 N 5 6 6 3 11 23 7
13.6 IX 17 13 15 15 1,3 18 23 N N N N N N 7
13.6* IX 17 13 14 14 1,3 11 N 4 6 6 3 12 22 7
Table 4.-
-Measurements (mm) of larvae of Artedius harringtoni. (Specimen between dashed line is undergoing noto-
chord flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length '
Parietal
spine length
3.0
4.7
0.66
0.82
0.16
0.20
0.32
0.39
1.3
1.6
0.72
1.1
0.12
0.31
N2
N
-
6.9
1.8
0.39
0.58
3.3
1.6
0.47
N
-
7.3
9.3
13.6
2.0
2.0
4.3
0.47
0.35
1.1
0.78
0.94
1.1
3.4
4.5
6.6
2.4
3.0
3.8
0.90
2.0
3.6
N
N
N
—
'Multiple spines develop along preopercular margin instead of usual 4.
5
2N = Not measured.
Artedius harringtoni
3.0mm NL
4.7mm NL
6.9mm NL
Figure 1. — Larvae of Artedius harringtoni.
Arfedius harringtoni
!
)
/'
7.3mm SL
9.3mm SL
13. 6mm SL
Figure 2. — Larvae of Artedius harringtoni.
ded in later stages, >9 mm. A row of >20 melanophores
(often 20-30 but >25 in larvae <6 mm) occurs along the
ventral midline beginning immediately behind the anus.
The number of ventral midline melanophores helps to
distinguish A. harringtoni from Artedius Type 2. The
posterior half of these ventral melanophores characteris-
tically extend as dashes of pigment out onto the finfold
particularly in larvae >7 mm. Some pigment is added to
the head region and pectoral fin base in the largest
specimens >12 mm.
The preopercular margin develops a series of small
spines which begin to appear in larvae ~4. 5-5.0 mm.
These increase in number to ~ 20-25 in postflexion lar-
vae. The spines on the dorsal edge and at the posterior
angle of the preopercle become the largest in the series.
The number of preopercular spines becomes reduced in
the largest specimens, possibly due to fusion or over-
growth by tissue. Artedius harringtoni never develops
pronounced multiple spines in the parietal and post-
temporal region as does Artedius Type 2 although mi-
nute serrations are sometimes visible and a bump
develops in the parietal region in larger specimens.
Larvae of A. harringtoni are stubby in shape and often
have a humped appearance in the nape region. The
hindgut trails below the body. Snout to anus length is
generally <50cc SL. Gut diverticula, or protrusions from
the abdominal wall, are never present as in Artedius
Type 2. Body depth at pectoral fin base ranges from
about 23 to 33co SL. The length of the pectoral fin in-
creases from 4 to 26co SL during larval development.
Artedius harringtoni is the only species of Artedius with
seven branchiostegal rays (Howe and Richardson foot-
note 3). Seven branchiostegals were countable in all lar-
val specimens examined (25) down to ~7 mm. The inner-
most pair of branchiostegals is extremely fine and small
in relation to the other six and may not be formed or
indistinguishable on unstained specimens under 7 mm.
The adult complement or dorsal, anal, pectoral, and
pelvic fin rays or ray elements can be counted in larvae
~10 mm.
Relative abundance of adults in the area of larval cap-
ture was also helpful. Artedius harringtoni is one of the
three most common species of Artedius in Oregon
tidepools, together with A. lateralis and A. fenestralis
(Howe6). Cirri patterns on the largest specimens in our
series compared with those known for these three species
provided further evidence of their identity as A. har-
ringtoni. Within these three species, only A. harringtoni
has preorbital cirri and only A. harringtoni and A.
lateralis have nasal cirri (Bolin 1944). Artedius har-
ringtoni has 1 small cirrus at the base of each nasal spine
while A. lateralis has 1 or 2 cirri at the base of each spine.
The largest specimens in our series have a single nasal
cirrus at the base of each spine and the largest specimen
has small bumps in the preorbital region which may be
developing cirri.
Artedius Type 2
(Figures .3, 4; Tables 5, 6)
Literature. Blackburn (1973) briefly described and
illustrated (8.5 mm SL) larvae resembling our Artedius
Discussion. Identification of this series as A. har-
ringtoni was based primarily on meristic characters, es-
pecially the presence of seven branchiostegal rays.
K. Howe, Ph.D. candidate, Department of Fisheries and Wildlife,
Oregon State University, Corvallis, OR 97331, pers. commun. September
1978.
Table 5. — Meristics from larvae of Artedius Type 2. (Specimen between dashed line is undergoing notochord flexion.)
[* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; B = Bases only; N = Not
examined.]
Anal Pectoral
Pelvic
fin
Preoper-
Ventral
midline
Caudal fin rays
Dorsal
Ventral
Vertebrae Branchi-
Body
length
Dorsal fin
Spines Rays
fin
rays
fin rays
Left Right
spine
& rays
cular
spines
melano-
phores
Second-
ary
Prin-
cipal
Prin-
cipal
Second-
ary
Abdom-
inal
Cau-
dal
ostegal
rays
3.0
4.7
— —
—
— N
— N
—
~5
24
14
N
X
N
N
N
X
N
X
N
N
N
N
N
X
6.0
- -
-
N
-
13
-15
N
N
N
N
N
N
N
7.3
-14B-**
12B
—
N
—
21
15
N
N
N
N
X
X
6
9.9
VIII 18
12
15
N
1,3
23
18
N
X
N
N
X
N
6
11.5*
IX 16
12
15
15
1,3
24
X
—
6
4
—
12
22
6
11.8
IX 17
13
15
N
1,3
13
13
X
N
N
N
N
N
6
Table (i.
-Measurements (mm) of larvae of Artedius Type 2. (Specimen between dashed line is undergoing notochord
flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length"
3.0
4.7
0.66
1.2
0.18
0.12
0.30
0.44
1.2
1.9
0.58
1.2
0.22
0.20
N3
X
N
X
6.0
1.7
0.35
0.56
2.8
1.5
0.31
N
N
7.2
9.9
11.8
2.0
3.0
4.1
0.31
0.48
0.64
0.66
0.94
1.13
3.5
4.6
5.6
2.1
2.8
3.0
0.86
1.7
3.4
N
N
N
N
N
N
"Multiple spines develop along preopercular margin instead of usual 4.
"Multiple spines develop in region of parietal spine.
3N = Not measured.
Artedius Type 2
3.0mm NL
4.7mm NL
6.0mm NL
Figure 3.-Larvae of Artedius Type 2. The second figure from the top is a dorsal view showing gut diverticula.
9
Artedius Type 2
7.2mm SL
1.8mm SL
Figure 4. — Larvae of Artedius Type 2.
Type 2 which he called Cottid 4, "Water Wings."
Eldridge (1970) also described and illustrated (3.2 mm)
similar appearing larvae with gut protrusions called Cot-
tid No. 4. White (1977) illustrated (3.9 mm NL) and
briefly described similar larvae as Cottid III with two
"horn-like extensions protruding . . . from the gut."
Young stages of two known species, A. lateralis and A.
fenestralis, have been described which somewhat resem-
ble our Artedius Type 2, although their similarity cannot
be confirmed on the basis of the literature alone (see
Discussion below). Budd (1940) illustrated and described
eggs and larvae (4.1 mm SL) of A. lateralis. Marliave
(1975) illustrated and briefly described young stages (4,
8, 11, 14 mm TL (total length)) of A. lateralis. He also
discussed eggs of A. lateralis and eggs and larvae of A.
fenestralis. Larvae of this form were called Artedius sp. 2
by Richardson (footnote 4) and Richardson and Pearcy
(1977).
Distinguishing Features. Larvae of Artedius Type 2
probably hatch at around 3-4 mm. The largest specimens
we have captured pelagically are ~14 mm, taken in a
neuston net 2 km off Newport, Oreg. They are beginning
to develop juvenile pigmentation.
10
In larvae, melanophores are concentrated over the
dorsolateral surface of the gut. Nape pigment is usually
present in larvae except at the smallest sizes but
becomes embedded and obscured in later stages >8 mm.
A row of usually <20 (often 15-20 but <25 in larvae <5
mm) melanophores occurs along the ventral body
midline posterior to the anus. The number of ventral
midline melanophores helps distinguish Artedius Type 2
from A. harringtoni which usually has >20. Approxi-
mately the posterior half of these melanophores extend
as characteristic dashes of pigment onto the ventral fin-
fold especially in larvae >7 mm. In the largest
specimens, ~12-14 mm, pigment is added to the head
region, at the base of the pectoral fin, to the anterior por-
tion of the spinous dorsal fin, as a bar extending from the
dorsal surface of the abdominal cavity near the dorsal
origin of the pectoral fin to the middle of the spinous dor-
sal fin, and as groups of melanophores along the lateral
midline from the gut region to the caudal fin.
As in A. harringtoni a series of small spines develops
along the margin of the preopercle, beginning to appear
~4.5-5.0 mm. They increase in number to ~ 20-25 in
postflexion larvae, — 7-11 mm, then decrease in the larg-
est specimens. Those at the dorsal edge and at the pos-
terior angle of the preopercle become large relative to the
rest of the spines. Clusters of spines develop in the
parietal region beginning with 1 or 2 spines in 6-7 mm
larvae, 3 spines in 7.5-8.5 mm larvae, and more on larger
specimens. By ~9-9.5 mm clusters of spines occur in
both the parietal and posttemporal regions, and these re-
main visible on the largest specimens examined. Such
spine clusters are never obvious on A. harringtoni.
Larvae of Artedius Type 2 are similar in shape to A.
harringtoni being stubby and often with a hump in the
nape region. The hindgut appears to trail below the
body. Snout to anus length is <50% SL. A prominent
feature is the presence of gut diverticula, one on each
side, protruding laterally from the dorsal surface of the
abdominal cavity immediately posterior to the pectoral
fin. These gut diverticula, never observed in A. har-
ringtoni, are most pronounced in small larvae, becoming
less conspicuous as the pectoral fin develops. They are
present on the largest pelagic specimens we examined.
This larval character is probably lost during transforma-
tion to the juvenile stage. Body depth at the pectoral fin
base ranges from about 19 to 29% SL. The length of the
pectoral fin increases from 4 to 29% SL during larval
development. Branchiostegal rays, which could be
counted on specimens >7 mm, consistently numbered 6
for all larvae in this series compared to 7 for A. har-
ringtoni. The adult complement of dorsal, anal, pectoral,
and pelvic fin rays or ray elements is countable by ~10
mm.
Discussion. Meristics on the largest specimens indicate
they may be either A. fenestralis or A. lateralis, two of
the three most abundant species of Artedius off Oregon,
as previously noted. Dorsal soft ray counts of 16-18
(16=6; 17=8, 18 = 2) on larval specimens in our collec-
tions seem more indicative of A. fenestralis than A.
lateralis which has 15-17 (Howe and Richardson footnote
3).
Head cirri patterns on the largest specimens indicate
they could be A. fenestralis. The largest specimens of
Artedius Type 2 have postocular cirri but show no sign of
forming nasal cirri. Bolin (1944) reported that A.
fenestralis has no nasal cirri while A. lateralis has 1 or 2
cirri at the base of each nasal spine. This would indicate
Artedius Type 2 is A. fenestralis unless nasal cirri appear
later in development. The largest specimens of A. har-
ringtoni, similar in size to the largest specimens of
Artedius Type 2, already have nasal cirri, indicating that
the absence of nasal cirri in Type 2 is real.
In a brief description Marliave (1975) stated that
". . . yolk sac larvae of Artedius fenestralis are identical
to the same stage of A. lateralis except that they are
perhaps a bit smaller. By the onset of fin ray formation
A. fenestralis larvae have a patch of melanophores on the
nape but none on the head, in contrast to A. lateralis . . .
[which] develops a patch of melanophores in the oc-
cipital region (none on the nape)." He further stated that
A. lateralis larvae "develop lateral extensions . . . over
the gut" that "distend the body wall over the dorsal in-
sertion of the pectorals." Presumably then both A.
lateralis and A. fenestralis larvae have gut diverticula.
[Budd (1940) did not show gut diverticula on larvae of A.
lateralis, but his specimens may have been too small to
have developed them.] However, if the presence of nape
pigment is a specific character for A. fenestralis as
Marliave reported, then most, if not all, of our Artedius
Type 2 may be A. fenestralis. That is if the other larval
characters of <20 ventral midline melanophores, pres-
ence of gut diverticula, and a pronounced cluster of
spines in the parietal region are adequate to distinguish
Artedius Type 2 at the species level. Interestingly, larvae
of this type occur off southern California (Ahlstrom7)
beyond the southern range limit of Diablo Cove, Calif.,
reported for A. fenestralis (Miller and Lea 1972).
However, A. lateralis does occur there. Final confirma-
tion of our Type 2 awaits identification of larvae of all
Artedius species reported to occur off Oregon.
If this series is A. fenestralis and we have also taken A.
harringtoni, we would expect to have larvae of A.
lateralis in our collections since adults of all three are
relatively abundant in Oregon tidepools, as we have
noted. Larvae of A. harringtoni and Artedius Type 2 were
among the most abundant cottid larvae in our collec-
tions. Explanations for missing A. lateralis larvae could
be several: 1) they are mixed in with one of our series,
e.g., Artedius Type 2; 2) they are one of our unidentified
larval types and not as abundant in our collections;
3) they exhibit a different early life history strategy, e.g.,
they are more benthic or coastal in nature and thus
would not be found in plankton collections taken 2 km or
more offshore.
E. H. Ahlstrom, Senior Scientist, Southwest Fisheries Center,
National Marine Fisheries Service, NOAA, P.O. Box 271, La Jolla, CA
92038, pers. commun. October 1978.
11
Clinocottus acuticeps (Gilbert)
(Figures 5, 6; Tables 7, 8)
Literature. An 8.6 mm SL larva resembling Clinocottus
acuticeps was illustrated and described by Blackburn
(1973). He called his specimen Cottid 1 "Biramous
anus," possibly Clinocottus sp. Eggs and larvae of two
Figure 5. — Larvae of Clinocottus acuticeps. The second figure from
the top is a ventral view showing hindgut diverticula.
Clinocottus acuticeps
40tti
3.7 mm NL
3.7mm NL
3.9mm NL
6.9mm NL
12
Clinocottus acuticeps
7.6mm SL
16.5mm SL
Figure 6. — Young of Clinocottus acuticeps.
13
other species of Clinocottus from the northeast Pacific
have been described, C. analis ([Eigenmann 1892 (as
Oligocottus analis); Budd 1940] including illustrations of
larvae and juveniles 4.6, 5.0, 7.6, 8.3, 9.9, 10.8, 18.0, and
24.3 mm TL. Larvae of C. acuticeps were called Cottidae
sp. 12 by Richardson (footnote 4) and Richardson and
Pearcy (1977).
Distinguishing Features. Larvae of C. acuticeps prob-
ably hatch at ~3-4 mm and begin to transform to
juveniles ~13-14 mm. A 16.5 mm specimen collected in a
tidepool is fully transformed juvenile.
Pigment is moderately scattered over the dorsolateral
surface of the gut. Nape pigment is present on all small
larvae but becomes embedded in larger specimens. Pig-
ment is also present on the head between the eyes in all
but the smallest specimens which may be faded. Melano-
phores are added to the head region during development.
A series of usually <15 ventral midline melanophores is
present on the tail beginning at about a vertical through
the anal opening in the trailing hindgut. No melano-
phores are on the ventral body margin anterior to that
point. Melanophores in this ventral midline series are
rather inconspicuous compared to other species. The pos-
terior melanophores in this ventral row do not extend as
dashes onto the finfold as in Artedius spp. but may ac-
tually occur on the ventral finfold rather than on the
body margin in larger larvae. A single melanophore per-
sists near the notochord tip, eventually appearing at the
middle of the caudal fin base. A few streaks of pigment
may be visible on the ventral finfold near the tail tip and
later on the caudal fin. On larvae nearing transforma-
tion, pigment is added to the head, pectoral fin base, and
onto the anterior end of the spinous dorsal fin. Recently
transformed juveniles are heavily pigmented with a
noticeably darkened area at the anterior end of the first
dorsal fin as in adults.
A series of ~10-15 small spines develops on the
preopercular margin. These begin to appear at ~6-7 mm.
With development, the upper spines become the most
prominent. The number of preopercular spines de-
creases during transformation with only 1 dorsal spine
visible in a 19.4 mm stained juvenile. No additional head
spines are obvious in larvae. Parietal and nuchal spines
are never apparent although the flabby skin could pos-
sibly obscure their presence on unstained larvae.
Larvae of C. acuticeps have an unusual gut, the poste-
rior portion of which trails well below the body. Near the
anus two diverticula or protrusions are evident through-
out the larval period, but they are no longer visible on
transformed juveniles. Snout to anus length is relatively
long for cottid larvae, ranging from ~62 to 71% SL during
the larval period. The larvae have a rather flabby
appearance at times seeming to be encased in a bubble of
loose outer skin, particularly in the head region. Body
depth ranges from ~22 to 33% SL during larval develop-
Table 7. — Meristics from young of Clinocottus acuticeps. (Specimen between dashed line is undergoing notochord flexion.)
(* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; B = Bases only; D =
Damaged; N = Not examined.]
Pelvic
Ventral
Caudal fin ravs
Dorsal fin
Anal
fin
Pectoral
fin rays
fin
spine
Preoper-
cular
midline
melano-
Dorsal
Ventral
Vertebrae
Abdom- Cau-
Branchi-
Body
Second-
Prin-
Prin-
Second-
ostegal
length
Spines Rays
rays
Left
Right
& rays
spines
phores
ary
cipal
cipal
ary
inal
dal
rays
3.7
— —
—
—
N
—
—
7
N
N
N
N
N
N
N
3.9
— —
—
—
N
—
—
9
N
N
N
N
N
N
N
6.9
-22B-**
11B
14
N
-
11
8
N
N
N
N
N
N
N *
7.6
VII 15
11
14
N
Buds
11
7
N
N
N
N
N
N
N
10.2*
VIII 15
12
14
14
1,3
11
N
4
6
5
2
10
14
6
10.4
VIII 16
12
1 1
N
1,3
13
7
N
N
N
N
N
N
N
13.8
IX 15
11
14
N
1,3
11
4
N
N
N
N
N
N
N
14.2*
IX 16
13
14
14
1,3
12
N
7
6
5
5
10
22
6
19.4*
D D
12
14
14
1,3
1
N
D
D
D
1)
10
23
6
Table 8. — Measurements (mm) of larvae of Clinocottus acuticeps. (Specimen between dashed line is undergoing noto-
chord flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length1
Parietal
spine length
3.7
3.9
0.90
0.88
0.14
0.16
0.38
0.38
2.3
2.4
0.90
0.86
0.26
0.40
N"
N
-
6.9
2.0
0.39
0.62
4.6
2.1
0.82
N
-
7.6
10.4
13.8
2.3
2.9
1 2
0.47
0.58
0.86
0.70
0.94
1 2
5.4
7.0
9.0
2.5
2.6
4.2
1.2
1.6
4.1
N
N
N
-
Multiple spines develop along preopercular margin instead of usual 4.
14
"N = Not measured.
ment. The adult complement of fin rays (or elements)
can be counted in larvae by ~7-8 mm. Late stage larvae
have relatively long pectoral fins, ~30% SL. On larger
larvae, ~13-14 mm, the inner pelvic fin ray is noticeably
attached to the belly by a membrane. This diagnostic
character helped link the larval series to identified
juveniles.
Discussion. Larvae of C. acuticeps are similar to
described larvae of C. analis and C. recaluus in size at
hatching, ~3-4 mm, and presence of nape and head pig-
ment and pigment over the gut on small larvae. (The
lack of head pigment on the smallest larvae of C.
acuticeps may have been caused by fading.) All three
species have a series of ventral midline melanophores but
those on C. analis and C. recalvus appear to be higher in
number than C. acuticeps. The posterior melanophores
in this series do not appear to extend out onto the finfold
in C. acuticeps as reported for the other two species.
Based on the illustrations, small larvae of C. analis and
C. recalvus have a much shorter relative snout to anus
length than C. acuticeps. Large larvae of C. recalvus
develop multiple spines on the preopercle as do C.
acuticeps, but they apparently do not develop the
hindgut diverticula or the flabby appearance of C.
acuticeps. These marked differences in body form, main-
ly the gut length and hindgut protrusions of C. acuticeps
could lead to questions about the relationships within
the genus Clinocottus, particularly as larvae of more
species become known.
Cottidae Type 1
(Figure 7; Tables 9, 10)
Literature. Larvae of this type have not been
previously described. These larvae were called Cottidae
sp. 1C by Richardson (footnote 4) and Richardson and
Pearcy (1977).
Distinguishing Features. The smallest larvae in our col-
lections are ~4 mm. They are relatively undeveloped and
probably recently hatched. Our largest specimens are ~9
mm. They have visible fin ray elements but fins are not
completely developed. Transformation occurs at some
larger size.
The larvae are relatively un pigmented. Melanophores
occur at the nape, in a rather light scattering over the
dorsolateral gut surface and in a row of <10 along the
ventral midline posterior to the anus. These ventral
midline melanophores are well spaced beginning ~4-5
myomeres behind the anus and extending to the poste-
riormost myomere. They decrease in number anteriorly
with development.
The preopercle develops a high number, ~19, of small
spines on its margin. A cluster of tiny spines develops in
the parietal region, a few of which are visible on 4 mm
specimens. Another cluster of minute spines develops in
the posttemporal-supracleithral region on larger
specimens.
The larvae are rather stubby in shape. The hindgut
trails below the body. Snout to anus length is ~ 40-50%
SL. Body depth at pectoral base is -23-26% SL. The
snout is relatively round in appearance.
Discussion. Specific or generic identification of Cot-
tidae Type 1 is not possible without additional material.
These larvae are related to the Artedius-Clinocottus-Oli-
gottus group of cottids based on counts of the largest
specimen and the presence of multiple spines on the
preopercle. Based on larval characters, these larvae ap-
pear to most closely resemble A rtedius spp. The clusters
of spines in the parietal and posttemporal-supra-
cleithral region in these larvae have been observed only
in Artedius Type 2 and Cottidae Type 3. They do not oc-
cur in C. acuticeps, C. recalvus (Morris 1951), or 0.
maculosus (Stein 1972, 1973) the latter of which
reportedly develops only 2 spines in the parietal region.
Spines on the preopercle of the largest specimen of Type
1 appear to be enlarging somewhat dorsally and at the
posterior angle as in A. harringtoni and Artedius Type 2.
However, their development would have to be traced in
Table 9. — Meristics of larvae of Cottidae Type 1. (Specimen be-
tween dashed lines is undergoing notochord flexion.)
Body
length
Dorsal
fin
spines
Dorsal
fin
rays
Anal
fin
rays
Pectoral
fin rays
Left Right
Pelvic fin
spine
and rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
4.2
-
-
-
- N1
-
-
7
6.7
-
—
—
— N
-
19
7
9.0
IX
18
12
15 N
Buds
19
3
'N = Not examined.
Table 10.— Measurements (mm) of larvae of Cottidae Type 1 . (Specimen between dashed lines is undergoing notochord
flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length1
Parietal
spine length-'
4.2
0.84
0.20
0.42
1.8
0.98
0.20
N3
N
6.7
1.4
0.20
0.52
2.6
1.6
0.55
N
N
9.0
2.1
0.25
0.94
4.2
2.5
1.2
N
N
Multiple spines develop along preopercular margin instead of usual 4.
Multiple spines develop in region of parietal spine.
N = Not measured.
15
Cottidae Type
4.2 mm NL
6.7mm NL
■-■\
S
9.0mm SL
Figure 7.— Larvae of Cottidae Type 1.
larger specimens to determine positively if the pattern is
the same as in our two Artedius types. This preopercular
spine pattern differs from that in C. acuticeps and C.
recalvus (Morris 1951) in which the dorsalmost spine
becomes the largest. The number of preopercular spines
in Type 1 is relatively high, ~ 19 as in A. harringtoni and
Artedius Type 2 (>20), compared with the lower
numbers for C. acuticeps (10-15), C. recalvus (4-9), and
0. maculosus (7-9). The gut shape and length resemble
our Artedius types.
Based on the meristic compilation by Howe and
Richardson (footnote 3) (Table 1), counts on our largest
Cottidae Type 1 specimen, particularly the 18 dorsal soft
rays, agree with only four species: A. fenestralis, A. har-
ringtoni, 0. maculosus, and 0. snyderi. The larvae are
not A. harringtoni based on our description and they are
not 0. maculosus based on the description by Stein
(1972, 1973). Identification as 0. snyderi is not possible
based on Stein's (1972) description although it seems un-
likely that our larvae are this species. Larvae of 0.
maculosus have a very different preopercular spine pat-
tern from our Type 1, and one would expect a pattern
similar to 0. maculosus on 0. snyderi. Problems in iden-
tifying A. fenestralis were discussed under Artedius Type
2 and cannot be resolved at this time.
L6
Cottidae Type 2
(Figure 8; Tables 11, 12)
Literature. Larvae of this type have not been
previously described. These larvae were called Cottidae
sp. 20 by Richardson (footnote 4) and Richardson and
Pearcy (1977).
Distinguishing Features. The smallest larvae in the
series are ~4 mm and recent);- hatched. The largest
specimen is 7.4 mm in which tb-j bases of dorsal and anal
Cottidae Type 2
6.4mm NL
7.4mm NL
Figure 8.— Larvae of Cottidae Type 2.
17
fin rays are just forming. Size at transformation is un-
known.
Pretlexion larvae have pigment at the nape, a rela-
tively light scattering of melanophores over the dorso-
lateral surface of the gut and a series of <20 but >10
ventral midline melanophores posterior to the anus. This
ventral series begins near or several myomeres posterior
to a vertical through the anal opening and extends to
about the fourth from last myomere. Some melano-
phores in this series radiate out onto the finfold. A few
melanophores occur on the finfold near the tail tip. With
development some melanophores are added to the head
and the number of ventral midline melanophores de-
creases. Melanophores were not apparent on the caudal
fin base of the largest specimen, as would be expected
based on the pigmentation near the tail tip of smaller lar-
vae, but the specimen was badly faded.
A series of approximately 10 small spines develop on
the preopercular margin. The upper spine appears some-
what thicker than the lower spines on the largest
specimen. A tiny parietal spine is visible on the 7.4 mm
specimen but no additional head region spines are ap-
parent.
The shape of the gut is rather distinctive with the
hindgut trailing noticeably below the body, similar to
Clinocottus acuticeps. Snout to anus length is around
50% SL, longer than most species. When viewed ven-
trally, the wall of the hindgut bulges out on each side of
the anus reminiscent of the hindgut diverticula of C.
acuticeps but not as pronounced. Body depth at the pec-
toral fin base is ~21-27% SL. The snout is quite rounded.
The larvae have a flabby appearance, particularly
around the head region, similar to C. acuticeps.
Discussion. The multiple preopercular spine pattern of
Cottidae Type 2 indicates this type is related to the Arte-
dius-Clinocottus-Oligocottus group. Based on a com-
bination of larval characters it seems to resemble
Table 11. — Meristics from larvae of Cottidae Type 2. (Specimens
below dashed line are undergoing notochord flexion.)
Dorsal
Body fin
length spines
Dorsal Anal Pectoral Pelvic fin
fin fin fin rays spine
rays rays Left Right and rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
4.0
15
10
11
17
6
Clinocottus most closely. It has a relatively low number
(<15) of multiple spines on the preopercle as does C.
acuticeps (10-15) and C. recalvus (5-9) (Morris 1951).
The upper preopercular spine appears to be enlarging
relative to the rest, as in C. acuticeps and C. recalvus.
Only a tiny parietal spine is visible on the largest
specimen. Two tiny spines appear in the parietal and
nuchal positions in C. recalvus but only for a brief period,
and none were observed in C. acuticeps, possibly ob-
scured by the flabby skin. Melanophores appear on the
nape and head of all C. recalvus and all but the smallest
C. acuticeps and Cottidae Type 2. The lack of pigment
on the head of our smallest specimens could be a result of
fading. Clinocottus recalvus larvae were described from
reared and freshly preserved material which often tends
to have increased numbers of and more pronounced
melanophores. The hindgut trails well below the body as
in C. acuticeps. Bulges appear in the hindgut wall on
each side of the anus similar to the more pronounced
diverticula of C. acuticeps. Such protuberances were not
noted for C. recalvus, perhaps on observational over-
sight. Some melanophores in the ventral midline series
radiate onto the finfold as in C. recalvus. This quality
was not noted in C. acuticeps but possibly could be
related to quality of preservation. Melanophores occur on
the finfold near the tail tip, often as streaks of pigment in
all three forms. These melanophores later appear on the
caudal fin in C. acuticeps and C. recalvus. They were not
visible on our largest Cottidae Type 2 specimen possibly
because it was faded.
Cottidae Type 3
(Figure 9; Tables 13, 14)
Literature. Larvae of this type have not been
previously described. They were listed by Richardson
(footnote 4) and Richardson and Pearcy (1977) as
Oligocottus sp. 1.
Distinguishing Features. The smallest larvae we have
taken are ~5 mm and relatively undeveloped. Our
largest specimen is 12.5 mm. Most fins appear fully
formed except for secondary caudal rays, but final trans-
formation must occur at a somewhat larger size.
These larvae have intense pigment over the head and
nape even in the smallest specimens. The head pigment
is more pronounced than in any of our other lightly
Table 12. — Measurements (mm) of larvae of Cottidae Type 2. (Specimens below dashed line are undergoing notochord
flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
4.0
0.8
0.16
0.42
2.2
0.84
0.24
N2
-
6.4
7.4
1.5
1.8
0.27
0.55
0.55
0.66
3.0
3.9
1.6
2.0
0.27
0.66
N
N
0.10
Multiple spines develop along preopercular margin instead of usual 4.
N = Not measured.
18
Cottidae Type 3
* -■* ** »'<* $ * *«\V_, •
7.5mm NL
12.5 mm SL
Figure 9. — Larvae of Cottidae Type 3.
pigmented forms, i.e., those with minimal body pigment.
Pigment over the dorsolateral surface of the gut is also in-
tense and densely concentrated, appearing as large round
melanophores. The only other pigment consists of ~4-5
ventral midline melanophores near the posteriormost
myomeres and a few at the base of the forming caudal
fin.
The preopercle develops a series of multiple spines
numbering ~ 15-20. The upper spine is the largest in the
12.5 mm specimen. A cluster of spines develops in the
parietal region, appearing as a double row. A similar
cluster develops in the posttemporal-supracleithral
region. A nasal spine is also apparent in the 12.5 mm
larva.
The hindgut is somewhat trailing and snout to anus
length is ~48-58% SL. The hindgut bulges slightly on
either side of the anus although this is not nearly as
pronounced as in Clinocottus acuticeps or Cottidae Type
2. Body depth at the pectoral fin base is ~ 22-29% SL.
The snout remains relatively rounded.
Discussion. Additional larger specimens are needed
before identification of Cottidae Type 3 can be ac-
complished with certainty. Based on current knowledge
19
of larval characters, particularly the multiple preoper-
cular spine pattern, it belongs with the Artedius-Clino-
cottus-Oligocottus group. The number of preopercular
spines is generally less than for A. harringtoni and
Artedius Type 2 which have >20, and greater than for C.
acuticeps, C. recalvus, and 0. maculosus which all have
<15 (Morris 1951; Stein 1973). The upper spine in Cot-
tidae Type 3 becomes the largest as in Clinocottus and
unlike Artedius. A cluster of spines develops in the
parietal region similar to Artedius Type 2 and Cottidae
Table 13.— Meristics from larvae of Cottidae Type 3. (Specimens
above dashed line are undergoing notochord flexion.)
Ventral
Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline
Body fin fin fin fin rays spine cular melano-
length spines rays rays Left Right and rays spines phores
6.3 - - - - - 7 5
7.5 — 19 4
12.5 IX 15 12 14 14 1,3 16 1
Table 14.-
-Measurements (mm) of larvae of Cottidae Type 3. (Specimens above dashed line are undergoing noto-
chord flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length1
Parietal
spine length'
6.3
7.5
1.4
1.9
0.31
0.58
0.62
0.74
3.0
4.2
1.4
2.2
0.27
0.56
N3
N
N
N
12.5
3.4
0.80
1.1
7.3
3.4
2.9
N
N
Multiple spines develop along preopercular margin instead of usual 4.
'Multiple spines develop in region of parietal spine.
' N = Not measured.
Type 1 and unlike C. acuticeps, C. recalvus, and O.
maculosus. However, spines in the parietal cluster ap-
pear to be fewer in number (~5) and occur in two rows in
contrast to Artedius Type 2 and Cottidae Type 1. A
cluster of ~4-5 spines also develops in the posttemporal-
supracleithral region somewhat like Artedius Type 2 and
Cottidae Type 1. The hindgut bulges slightly around the
anus similar to but much less pronounced than in C.
acuticeps and Cottidae Type 2. The hindgut trails more
than in Artedius but less than in C. acuticeps and Cot-
tidae Type 2. Cottidae Type 3 has fewer ventral midline
melanophores than Artedius or Clinocottus, and these
appear only along the posterior portion of the tail as
reported for O. snyderi (Stein 1972).
Meristics of the largest Type 3 specimen fit a number
of species of Artedius and Clinocottus as well as Oligo-
cottus maculosus (Table 1). Our specimens differ from
larvae of the latter species described by Stein (1972,
1973) in having 1) ~19 preopercular spines instead of 7-9,
2) multiple spines in the parietal region instead of 2, and
3) only 4-5 ventral midline melanophores instead of 11-
20. Thus they must be either an Artedius or a Clino-
cottus species.
Triglops sp.
(Figure 10; Tables 15, 16)
Literature. Larvae identified as Triglops sp., which
resemble our larvae, were illustrated (8.3, 12 mm SL) by
Blackburn (1973) but no descriptive information was
given. Larvae of other species of Triglops which have
been described include 10-18 mm T. pingeli (Ehren-
baum 1905-1909; Koefoed 1907 [cited by Khan 1972];
Rass 1949) from the Atlantic Ocean and Barents Sea and
8.4, 11.6, 18.9, 23.4 mm TL T. murrayi (Khan 1972) from
the Atlantic Ocean.
Distinguishing Features. Our smallest preflexion larva
is 6.9 mm and relatively undeveloped. The largest
Table 15. — Meristics from larvae of Triglops sp. (Specimens above
dashed line are preflexion, that below is postflexion.)
Ventral
Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline
Body fin fin fin fin rays spine cular melano-
length spines rays rays Left Right and rays spines phores
6.9 ————— — —
7.4 _ _ _ _ _ _ _
15.4 X 31 30 16 16 1,3 4+4
Table 16. — Measurements (mm) of larvae of Triglops sp. (Specimens above dashed line are preflexion, that below is
postflexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
6.9
7.4
1.9
1.8
0.20
0.20
0.90
0.86
2.6
3.1
1.5
1.4
0.39
0.39
—
-
15.4
4.4
0.80
1.4
6.8
3.2
2.9
0.62
BR1
'BR = Broken.
20
Trig lops sp.
6.9mm NL
15.4mm SL
Figure 10. — Larvae of Triglops sp.
specimen we have is 15.4 mm. It has a large complement
of fin rays or elements although the fins do not appear to
be fully formed and transformation probably takes place
at a somewhat larger size. We have taken only four
specimens.
The larvae have a distinctive, heavily pigmented gut,
and postanal ventral midline melanophores are absent.
Pigment occurs over the head and nape region of the
smallest larvae, but the nape pigment is no longer visible
on the largest specimen. Pigment also occurs on the pec-
toral fin base. The rest of the body remains un-
pigmented.
Head spines on the largest specimen include a double
row (anterior and posterior series) of 4 preopercular
spines, a parietal spine, a nuchal spine, 2 spines in the
posttemporal-supracleithral region, and 2 spines in the
postocular region.
The larvae are relatively slender with depth at the pec-
toral base ranging from 19 to 22% SL. Snout to anus
length ranges from 38 to 44% SL. The larvae have a high
number of myomeres, ~ 45-46, which distinguishes them
from most other cottids in the region. The oblique
mouth, angular jaw, and pointed snout further dis-
tinguish these larvae.
Discussion. No species of Triglops are reported from
Oregon waters although our Triglops sp. larvae were all
collected 28 km off the Oregon coast in March and April.
The larvae could be a product of spawning in more north-
ern waters off Washington where two species of Triglops
reportedly occur, T. macellus and T. pingeli (Hart 1973).
But coastal currents generally flow northward during
that time of year, minimizing the likelihood that the lar-
vae were transported southward. The occurrence of
relatively undeveloped larvae off the mid-Oregon coast
indicates a species of Triglops may occur and spawn off
Oregon.
Although we cannot be positive that total adult com-
plements of fin ray elements are formed in the 15.4 mm
specimen, the counts of dorsal and anal fin rays, 31 and
30 respectively, are too high for T. pingeli [maximum of
28 in both fins based on Howe and Richardson (footnote
3)1. The counts are within the range given for T. macellus
(27-31 in both fins) as are the pectoral fin ray counts of 16
(range 15-17, based on Howe and Richardson footnote 3).
However, Howe and Richardson listed a count of 51
vertebrae for one specimen of T. macellus. We are able to
count only 47 myomeres on our larvae, seemingly too low
for T. macellus. The range of variability of vertebrae
numbers is not yet known for T. macellus. If the one
count of 51 is at the high end of the scale, 4 less vertebrae
may be within the range of variability and our larvae
could well be T. macellus. This remains to be confirmed.
Alternatively, an undescribed species of Triglops occurs
off the Oregon coast which seems less plausible.
Larvae of T. pingeli from the Atlantic Ocean and
Barents Sea presumably have a row of postanal ventral
midline melanophores based on the illustrations by
Ehrenbaum (1905-1909) and Rass (1949) and the discus-
sion by Khan (1972) who compared them to larvae of T.
murrayi which also have postanal ventral midline
melanophores. In addition, T. pingeli also develops a row
21
of melanophores along the base of the second dorsal fin
by the size of 10 mm and a mediolateral row by 11 mm
[Koefoed 1907 (cited by Khan 1972)]. If larvae of T.
pingeli from the Pacific develop the same pigment pat-
tern as those from the Atlantic and Barent Sea, they
would be easily distinguishable from our Triglops sp. lar-
vae. Head and trunk pigment and shape of T. pingeli
resemble our Triglops sp. indicating a generic similarity.
Chitonotus pugetensis (Steindachner)
(Figures 11, 12; Tables 17, 18)
Literature. Larvae of this species have not been
previously described.
Figure 11. — Larvae of Chitonotus pugetensis.
Chitonotus pugetensis
3.0mm NL
6.3mm NL
8.5mm SL
22
Chitonotus pugetensis
II. 5mm SL
5.4mm SL
16. 6mm SL
Figure 12. — Young of Chitonotus pugetensis.
Table 17. — Meristics from larvae of Chitonotus pugetensis. (Speci-
mens above dashed line are preflexion, those below are postflexion.)
Ventral
Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline
Body fin fin fin fin rays spine cular melano-
length spines rays rays Left Right and rays spines phores
3.0 — 45
6.3 — 41
8.5 -15-' 14 — — 4 37
11.5 IX 16 16 16 16 4 24
15.4 X 16 16 18 18 1,3 4 26
16.6 X 15 15 17 17 1,3 4 24
'Dorsal fin spines and soft rays cannot be distinguished.
Distinguishing Features. The smallest larvae we have
taken are ~3 mm and probably recently hatched. The
largest pelagic specimen, 16.6 mm, is beginning to trans-
form to the juvenile stage as evidenced by increasing
pigmentation.
The dorsolateral surface of the gut is only moderately
pigmented. Additional pigment distinctively lines the
ventral margin of the abdominal cavity. Pigment occurs
on the head by 8.5 mm and is added to the head region
through later development. A series of melanophores
lines the ventral body margin postanally. These ventral
midline melanophores always number >20 and usually
>40 in preflexion larvae <6 mm. This is the highest
number of ventral midline melanophores for preflexion
23
Table IS
— Measurements (mn
0 of larvae
of Chitonotus pugetensis. (Specimens above dashed line are
below are postflexion.)
preflexion, those
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
3.0
6.3
0.66
1.7
0.23
0.43
0.70
0.62
0.54
2.6
1.1
1.5
0.20
0.31
-
—
8.5
11.5
15.4
16.6
2.2
3.5
4.6
6.2
0.72
0.88
1.1
1.8
0.80
1.1
1.6
1.7
4.0
5.3
7.4
7.6
2.0
2.8
3.8
3.9
0.64
16
2.6
4.0
0.12
0.23
0.31
0.40
0.04
BR'
0.12
'BR =
Broken.
larvae of any species described in this guide. The poste-
rior melanophores in this series appear to be closer
together than the anterior ones. The number and spacing
of the ventral midline melanophores help distinguish
small larvae. Melanophores occur along the base of the
caudal fin after it develops. As transformation to the
juvenile stage approaches pigment is added to the ante-
rior portion of the spinous dorsal fin, in the nape region,
in blotches along the lateral midline, and on the pectoral
fin.
The preopercle becomes armed with 4 prominent
spines, some of which are visible on larvae as small as ~6
mm. With development, the upper 2 spines become
larger than the 2 lower ones. Small parietal and nuchal
spines develop during the postflexion period and fuse
together as 1 spine. Several small spines also develop in
the posttemporal and supracleithral region. These latter
two sets of spines are inconspicuous on the 16.6 mm
specimen. A nasal spine is visible by 15 mm.
Small larvae of C. pugetensis somewhat resemble
liparid larvae in shape except that they have fewer
myomeres. With development, the snout becomes rather
pointed in appearance and the body shape resembles
that of Ice linus spp. where the body outline narrows dis-
tinctively near the caudal peduncle. Body depth at the
pectoral fin base remains ~ 23-25% SL. Snout to anus
length ranges from ~41 to 48% SL and pectoral fin length
increases to ~24% SL during the larval period. The adult
complement of fin rays (except dorsal spines) can be
counted in 11.5 mm larvae. The deep emargination be-
tween the third and fourth dorsal fin spines, a diagnostic
character for C. pugetensis, is noticeable by 11.5 mm.
Discussion. Chitonotus is a monotypic genus in the
northeast Pacific. The larvae are rather distinctive, and
with the aid of the above characters, they are not easily
confused with other forms off Oregon.
Cottus asper Richardson
(Figure 13; Tables 19, 20)
Literature. Larvae of Cottus asper were briefly de-
scribed and illustrated (5.5, 9.0, 10.8 mm TL) by Stein
(1972). We did not review literature on larvae of strictly
freshwater species of Cottus.
Distinguishing Features. Larvae hatch ~5 mm and
begin to develop juvenile pigmentation ~10 mm as evi-
denced by an increase in the number of melanophores in
the head region.
Cottus asper larvae are relatively lightly pigmented. A
few melanophores cover the dorsal surface of the gut and
several elongate melanophores line the ventral margin of
the abdominal cavity and throat. About 15-20 evenly
spaced melanophores occur along the ventral midline of
the tail beginning near the anus. These decrease in
number during development while a few melanophores
are added laterally over the gut and one appears at the
ventral edge of the preopercle. As transformation begins
more pigment is added to the head, base of pectoral fin,
over the gut, and along the base of the caudal fin.
Table 19. — Meristics from larvae of Cottus asper. (Specimen above
dashed line is preflexion, those below are postflexion.)
Body
length
Dorsal Dorsal
fin fin
spines rays
Anal
fin
rays
Pectoral Pelvic fin
fin rays spine
Left Right and rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
5.2
- -
-
-
N1 -
-
17
8.2
9.9
-19-'
IX 18
16B3
16
19
N
18
4
4
14
8
N = Not examined.
"Dorsal fin spines and soft rays cannot be distinguished.
' B = Bases only.
Table 20. — Measurements (mm) of larvae of Cottus asper. (Specimen above dashed line is preflexion, those below are
postflexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
5.2
0.9
0.12
0.36
2.1
0.78
0.51
-
-
8.2
9.9
1.9
2.6
0.35
0.66
0.55
0.74
3.8
4.2
1.4
1.8
0.62
2.5
-0.03
0.33
-
24
Cottus osper
5.2mm NL
8.2mm SL
9.9mm SL
Figure 13. — Larvae of Cottus asper.
Four pronounced spines develop on the preopercular
margin, evident in 8 mm larvae. No additional head
spines form with development.
The shape of C. asper larvae is rather distinctive, being
somewhat blennioid in appearance. This is partly related
to the characteristic coiling and shape of the gut which is
unlike any of the other cottids described here. Snout to
anus length is usually between 40 and 46% SL. Body
depth at the pectoral fin base is slender relative to body
length, usually <20% SL. Total fin rays or elements are
countable by ~10 mm.
Discussion. Two species of Cottus, C. asper and C.
aleuticus, are found in brackish water off Oregon. Only
larvae of C. asper have been identified in coastal plank-
ton collections. Ringstad8 provided evidence that in a
British Columbia coastal stream most C. asper spawn in
the estuary while C. aleuticus spawn primarily in fresh
water. Thus larvae of C. aleuticus may simply not occur
"Ringstad, N. R. 1974. Food competition between freshwater
sculpins (Genus Cottus) and juvenile coho salmon {Oncorhynchus ki-
sutch): an experimental and ecological study in a British Columbia
coastal stream. Environ. Can., Fish. Mar. Serv., Tech. Rep. 457, 88 p.
in coastal plankton. Larvae of C. aleuticus have not been
described. Meristics, especially anal (usually >15 for
asper and <15 for aleuticus) fin ray numbers will gen-
erally separate the two (Scott and Crossman 1973; Howe
and Richardson footnote 3).
Enophrys bison (Girard)
(Figure 14; Tables 21, 22)
Literature. Blackburn (1973) illustrated and briefly de-
Table 21. — Meristics from larvae of Enophrys bison. (Specimen be-
tween dashed lines is undergoing notochord flexion.)
Body
length
Dorsal
fin
spines
Dorsal
fin
rays
Anal Pectoral
fin fin rays
rays Left Right
Pelvic fin
spine
and rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
4.8
-
-
- - N'
-
-
14
7.0
-
-
— — N
—
1
. 11
9.1
IX
11
9 16 N
[,'i
4
10
N = Not examined.
2.r>
Enophys bison
4.8mm NL
7.0mm NL
9.1mm SL
Figure 14. — Young of Enophrys bison.
26
Table 22. — Measurements (mm) of larvae of Enophrys bison. (Specimen between dashed line is undergoing notochord
flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
4.8
1.1
0.2
0.55
2.3
1.1
0.37
-
-
7.0
2.0
0.5
0.66
3.8
1.9
0.55
0.35
0.27
9.1
3.4
0.6
0.90
5.0
3.4
2.08
0.43
0.27
scribed a 7.5 mm larva of E. bison and Marliave (1975) a
10 mm TL larva. Misitano (1978) described a develop-
mental series, including eggs, of E. bison (5.0, 5.4, 5.8,
6.7, 7.1, 7.6 mm SL) based on laboratory-reared
material.
Distinguishing Features. Larvae are ~5 mm at
hatching. The addition of juvenile pigment may begin in
specimens as small as 7 mm in the laboratory but may
occur later in the natural environment based on our
material. Transition from pelagic to benthic habitat
begins in specimens as small as 7.6-7.8 mm in the labo-
ratory but Marliave (1975) captured a 10 mm TL trans-
forming specimen in the plankton and we took trans-
forming specimens up to ~9.5 mm in the neuston. Even
with this variation, transformation to juveniles takes
place at a relatively small size in E. bison compared to
other known cottids.
Pigmentation over the dorsolateral surface of the gut is
characteristic and intense. Melanophores often appear as
tightly packed discrete circles. Additional melanophores
are on the ventrolateral surface of the gut of fresh mate-
rial but are faded on our smallest specimens. Melano-
phores are present on the nape, appearing very dark in
freshly preserved specimens. The number of postanal
ventral midline melanophores is usually <15. A dis-
tinctive space of several myomeres in length exists
between the anus and the anteriormost ventral melano-
phore. The ventral midline melanophore series is rela-
tively inconspicuous, compared to other cottid species.
During development, pigment is added to the head
which becomes entirely covered with melanophores by
transformation. Melanophores are added beneath the
spinous dorsal fin forming a band extending to the gut, to
the base of the pectoral fin, to the ventrolateral region of
the gut, and eventually beneath the second dorsal fin ex-
tending as a band to the ventral body margin. A few
melanophores appear at the base of the caudal fin.
Larvae of E. bison develop very prominent head
spines. Four pronounced preopercular spines are visible
on all but the smallest larvae. The length of the second
preopercular spine is 18% HL in a specimen undergoing
notochord flexion. They also develop a rather large
parietal spine, visible by ~6 mm. With development, a
nuchal spine appears posterior to the parietal and fuses
with it, and another spine develops on the parietal ridge
anterior to the parietal spine. A postocular spine
develops over the eye. A spinelike bump develops in late
larvae in the posttemporal-supracleithral region and a
nasal spine appears on the snout. Two spiny tips (mid-
opercular and interopercular-subopercular) can be seen
on the gill cover margin.
All but the smallest larvae are relatively deep bodied
with body depth at the pectoral fin base increasing to
~37% SL on our transforming specimens. Snout to anus
length increases from ~48 to 55% SL with development.
The preanal finfold described by Misitano (1978) in lar-
vae <6.6 mm was not visible on our specimens probably
because of poor condition. All fin rays are countable by
the onset of the transformation period at ~9 mm in our
material. Pectoral fin length reaches 23% SL by trans-
formation. Enophrys bison larvae have a character-
istically low number of myomeres (vertebrae = 29-31)
(Howe and Richardson footnote 3) which helps to distin-
guish them from other cottids off Oregon most of which
have >31 vertebrae (Table 1).
Discussion. Only one species of Enophrys occurs off
Oregon and the larvae are rather distinctive and not like-
ly to be confused with other species. However, two or
three additional species of Enophrys may occur in the
northeast Pacific, E. diceraus, E. lucasi, and E. taurina
(Sandercock and Wilimovsky 1968; Quast and Hall
1972). Their larvae have not yet been described. Whether
larval characters useful for E. bison (i.e., distinctive gut
pigment, intense nape pigment, pronounced head spines,
gap between anus and first ventral midline melano-
phore, small size at transformation) would be helpful in
distinguishing larvae of the other species remains to be
determined. Certainly the low number of vertebrae (26-
29) of E. taurina (Howe and Richardson footnote 3) will
be a useful character. Larvae of E. bubalis and E. lillje-
borgi (Russell 1976) closely resemble E. bison with some
differences in pigmentation and spination.
Icelinus spp.
(Figures 15, 16, 17; Tables 23, 24)
Literature. A 4.3 mm larva resembling this type was
illustrated and described by Blackburn (1973) as Cottid
3. These larvae were referred to as Icelinus sp. 1 by Rich-
ardson (footnote 4) and Richardson and Pearcy (1977).
Distinguishing Features. The smallest larvae in our col-
lections are ~3-4 mm and probably recently hatched.
The largest specimens we have collected pelagically are
27
Icelinus spp.
3.3mm NL
8.6mm NL
V— ^o..
* >:
10.9mm SL
Figure 15. — Larvae of Icelinus spp.
~ 17-18 mm, some taken in neuston tows, and they are
beginning to develop juvenile pigmentation.
Larvae up to ~8-9 mm are characterized by a series of
large melanistic spots on the dorsal and ventral finfold,
when intact. Most all of our specimens have 2 spots dor-
sally and ventrally but one had 3 ventral spots, and
Blackburn's (1973) specimen had 3 spots dorsally and
ventrally. Whether the number of finfold spots is a
specific character or merely individual variation is not
known at this time. Moderate pigment occurs over the
dorsolateral surface of the gut and some melanophores
are present near the base of the cleithrum. A series of dis-
tinctive melanophores, usually <15, occurs along the
ventral midline of the tail beginning immediately behind
the anus. These melanophores vary in size with some of
them appearing more pronounced than others. One or
several melanophores clustered at the posterior end of
this ventral row distinctively appear at the ventral
margin of the caudal fin base after it is formed. With
development pigment is added to the head and over the
gut with little additional change. Some larger speci-
mens, >12 mm, have 2 distinct melanophores on the dor-
sal midline in the tail region. Whether this is a specific
character is unknown, but we have been unable to link a
complete developmental series together based on the
presence of these dorsal melanophores. Some speci-
mens, with and without dorsal midline melanophores,
have an additional distinct pigment spot on the dorsal-
most rays of the caudal fin near its base. In specimens
nearing transformation, pigment increases over the head
region, on the spinous dorsal fin, and on the pectoral fin
base.
28
Icelinus spp.
13. 8mm SL
15. 2mm SL
6.5mm SL
Figure 16. — Young of Icelinus spp.
Spines are first noticeable on the preopercle ~8-9 mm
and 4 are obvious by 10-11 mm. The upper one becomes
somewhat larger than the rest. Parietal and nuchal
spines develop and fuse together. Several spines appear
in the posttemporal-supracleithral region eventually be-
ing reduced to bumps. These are difficult to distinguish
from developing spinelike scales along the lateral line
and below the spinous dorsal fin. A nasal spine is obvious
on the largest specimens.
The gut shape is rather distinctive in larvae of Ice-
linus spp. Snout to anus length is relatively short in
small larvae, <40% SL, but increases to ~50% SL on
larger specimens. Body depth at the pectoral fin base is
-20% SL in small larvae and -25-30% SL on older lar-
vae. Larvae of Icelinus spp. have a very characteristic
body shape. Body depth appears to be constricted just
behind the anus, bulges out slightly in the midtail region,
and then narrows distinctively again near the tail tip or
caudal peduncle. They also develop a rather pointed
snout which is apparent on larvae >8 mm. Pectoral fin
length is ~ 25-30% SL on the largest specimens. The
adult complement of fin rays or elements is countable on
specimens >12 mm including the diagnostic pelvic fin
count of 1,2.
Discussion. Identification of the largest specimens to
the genus level was relatively easy since Icelinus is the
only representative off Oregon except Zesticelus pro-
fundorum with 2 pelvic fin rays. The distinctive body
shape and pigmentation helped link the larval series to-
29
Icelinus spp.
^ ^^^^
12. 5mm SL
16.6mm SL
Figure 17. — Young of Icelinus spp.
Table 23. — Meristics from larvae of Icelinus spp. (Specimen between dashed line is undergoing notochord flexion.)
[* = Stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; t = Form with two melanophores on
dorsal midline; N = Not examined.]
Pelvic Ventral Caudal fin rays
Anal Pectoral fin Preoper- midline _ Dorsal Ventral Vertebrae Branchi-
Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal
length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays
3.3 10NNNNNNN
5.1 11 N N N N N N N
8.6 3 10 N N N N N N N
10.9 -25-** 12 15 15 4 10 N N N N N N N
tl2.5 X 16 12 16 16 1,2 4 8 N N N N N N N
13.8 X 16 12 16 16 1,2 4 10 N N N N N N N
1 14.1 X 16 12 16 16 1,2 4 11 N N N N N N N
14.4* XI 15 12 15 15 1,2 4 N 5 6 6 4 11 23 6
15.2 X 16 12 15 15 1,2 4 11 N N N N N N N
U5.5* X 16 12 15 15 1,2 4 N 6 6 6 4 11 23 6
16.5 X 16 12 16 15 1,2 4 10 N N N N N N N
U6.6 X 16 13 15 15 1,2 4 12 N N N N N N N
gether at least at the generic level. However, specific
identification is complicated because most meristics of
the four Icelinus species reported to occur off Oregon, /.
burchami, I. filamentosus, I. oculatus, and /. tenuis
overlap. An anal fin ray count of 12 agrees only with /.
burchami from Oregon waters (Table 1), a species which
is reportedly rare (Howe and Richardson footnote 3).
Four other northeast Pacific species of Icelinus (I.
borealis, I. cavifrons, I. fimbriatus, and /. quadri-
seriatus) can have an anal ray count of 12 but none
reportedly occur off Oregon (Howe and Richardson foot-
note 3). Characters used to distinguish adults such as cir-
ri patterns and elongated dorsal fin spines are not
developed on our largest pelagic specimens. These fac-
tors together with inability to find consistent larval char-
acters to split our Icelinus spp. larvae into subgroups po-
tentially equivalent to species forces us to keep our
indentification at the generic level.
30
Table 24. — Measurements (mm) of larvae of Icelinus spp. (Specimen between dashed line is undergoing notochord
flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
3.3
5.1
0.76
0.86
0.10
0.12
0.32
0.39
1.2
1.7
0.68
1.0
0.26
0.31
-
—
8.6
1.8
0.51
0.55
3.4
1.7
0.39
-
-
10.9
n.2.5
13.8
U4.1
15.2
16.5
16.6
2.5
4.0
4.2
5.2
5.2
7.2
6.1
0.80
1.0
1.0
1.4
1.2
2.7
1.8
0.88
1.2
1.3
1.4
1.5
1.4
1.4
4.6
6.2
6.4
7.0
7.7
9.0
8.3
2.2
3.0
3.6
4.2
4.0
4.8
4.2
1.04
2.2
2.4
3.0
4.1
3.8
3.2
0.22
0.32
0.55
0.48
0.56
0.56
0.32
0.12
0.26
0.31
0.24
0.24
0.24
0.32
+Form with 2 melanophores on dorsal midline.
Leptocottus armatus Girard
(Figure 18; Tables 25, 26)
Literature. Eggs and recently hatched larvae (~4 mm
TL) were illustrated and described by Jones (1962). Eggs
and young (8, 12, 13 mm TL) were briefly discussed by
Marliave (1975) and larvae by Blackburn (1973). White
Figure 18. — Larvae of Leptocottus armatus.
Leptocottus armatus
5.1mm NL
i:;::y
8.1mm NL
ll.lmm SL
31
Table 25. — Meristics from larvae of Leptocottus armatus. (Specimen
between dashed line is undergoing notochord flexion.)
Body
length
Dorsal
fin
spines
Dorsal
fin
rays
Anal
fin
rays
Pectoral
fin ravs
Left Right
Pelvic fin
spine
and rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
5.1
-
-
-
-
N1
-
-
11
8.1
—
—
15
-
N
-
4
9
11.1
VIII
18
17
19
N
—
4
7
Discussion. Leptocottus is a monotypic genus in the
northwest Pacific. The larvae with their characteristic
bars of pigment over the gut should not be easily con-
fused with any other species unless damage to the speci-
men results in a blurring of the gut pigment. Even so, the
internal pigment on the snout serves as a useful
diagnostic character.
N = Not examined.
Table 26. — Measurements (mm) of larvae of Leptocottus armatus. (Specimen between dashed lines is undergoing
notochord flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
5.1
0.92
0.18
0.25
2.0
0.84
0.23
-
-
8.1
1.9
0.31
0.32
3.2
1.6
0.78
0.10
-
11.1
2.8
0.62
0.82
4.9
2.3
2.2
0.21
—
(1977) illustrated (4.9 mm NL) and briefly described
small larvae. Blackburn's Cottid 5 (5.8 mm SL) was
probably this species as was Eldridge's (1970) Cottid No.
I (5.1 mm).
Distinguishing Features. Hatching takes place at 3.9-
4.8 mm TL. Our largest pelagic specimen is 11 mm and
may be nearing transformation as indicated by in-
creased pigmentation over the head. Marliave's (1975) 13
mm TL specimen was transformed and had assumed a
benthic habit.
Larvae have distinctive pigment over the dorsolateral
surface of the gut which always appears as ~6-8 bars ex-
cept in damaged specimens. No other known cottid lar-
vae have such gut pigment. Larvae also have distinctive
internal pigment on the snout at the anterior part of the
forebrain and extending posteriorly from the eye to the
pectoral fin. The latter becomes obscured by muscle but
the former remains visible throughout the larval period.
Several melanophores occur along the ventral margin of
the gut cavity. A row of usually <15 rather prominent
melanophores occurs along the ventral midline of the tail
beginning about 5 myomeres posterior to the anus. With
development, melanophores are added to the head and a
few are present at the base of the caudal fin.
Four prominent spines develop along the preopercular
margin, evident in ~7 mm larvae. No additional head
spines are formed.
Larvae are rather slender with body depth at pectoral
fin base comprising 21% SL in the largest specimen.
Snout to anus length increases slightly from 39 to 44% SL
in the specimens examined. The snout has a rounded
appearance never becoming pointed. The adult comple-
ment of dorsal, anal, and pectoral fin rays is countable by
II mm; however, the pelvic fins are not developed at this
time.
Paricelinus hopliticus Eigenmann
and Eigenmann
(Figures 19, 20; Tables 27, 28)
Literature. Larvae have not previously been described.
Distinguishing Features. The smallest larvae in our col-
lections, ~5-6 mm appear to be recently hatched. An
18.6 mm specimen is beginning to transform as evi-
denced by increased head pigmentation and body spi-
nation (modified scales). A 25.6 mm specimen captured
pelagically appears to be a fully transformed juvenile.
Larvae have a distinctively pigmented gut with mela-
nophores covering the entire abdominal cavity. Pigment
is scattered over the head region. Over 30 melanophores
line the ventral body midline in the tail region beginning
just behind the anus in small larvae but the number de-
creases to ~ 15-20 in postflexion stages. At the posterior
end of this ventral row a separate group of distinctive
melanophores occurs near the tail tip extending onto the
finfold. These melanophores line the base of the caudal
fin after it forms. Little pigment is added during the lar-
Table 27. — Meristics from young of Paricelinus hopliticus. (Speci-
mens above dashed line are preflexion, those below are postflexion.)
Ventral
Dorsal
Dorsal
Anal
Pectoral
Pelvic fin
Preoper-
midline
Body
fin
fin
fin
fin
rays
spine
cular
melano-
length
spines
rays
rays
Left
Right
and rays
spines
phores
5.6
—
—
—
—
N1
—
—
32
6.2
—
—
—
—
N
—
—
31
13.8
XII
19
23
15
N
1,5
4
17
18.6
XII
19
23
15
N
1,5
4
17
25.6
XII
19
23
15
N
1,5
4
19
'N = Not examined.
32
Table 28. — Measurements (mm) of young of Paricelinus hopliticus. (Specimens above dashed line are preflexion, those
below are postflexion.)
Body depth
2d preoper-
Body
Head
Snout
Eye
Snout to
at pectoral
Pectoral
cular
Parietal
length
length
length
diameter
anus length
fin base
fin length
spine length
spine length
5.6
1.2
0.23
0.51
2.0
0.86
0.16
—
—
6.2
1.7
0.31
0.70
2.2
1.0
0.23
—
—
13.8
4.3
1.0
1.7
7.0
4.2
BR1
BR
BR
18.6
5.0
0.96
1.4
8.8
4.6
4.2
0.80
-0.24
25.6
8.6
2.1
2.7
11.5
5.3
7.4
0.96
0.48
'BR = Broken.
val period except over the head and at the base of the
pectoral fin. The pelagically captured juvenile has much
increased pigmentation in blotches over the entire body
and on most fins.
Four spines develop on the preopercular margin in
postflexion larvae. A postocular and a nasal spine also
become prominent. Spines appear to develop in the
parietal and nuchal region and also anteriorly along the
parietal ridge. These are difficult to distinguish from the
spiny scales that develop dorsally. Similarly, several
spines appear to develop in the posttemporal-supra-
cleithral region but are difficult to distinguish from the
spiny lateral line scales. Three spine tips (midopercular,
subopercular, interopercular) develop along the gill cover
margin. A cleithral spine becomes evident in the larger
specimens.
Snout to anus length is relatively short in preflexion
larvae, ~35-36% SL but increases with development.
Body depth at the pectoral base is also small relative to
body length, ~ 15-16% SL on preflexion larvae, in-
Paricelinus hopliticus
5.6mm NL
6.2mm NL
;s>
13. 8mm SL
Figure 19. — Larvae of Paricelinus hopliticus.
33
Paricelinus hopliticus
18. 6mm SL
25.6mm SL
Figure 20. — Young of Paricelinus hopliticus.
creasing to ~ 25-30% SL during the larval period then de-
creasing in the juvenile stage. The general body shape of
the larvae resembles that of Icelinus spp. The snout be-
comes noticeably pointed.
Discussion. Paricelinus is a monotypic genus in the
northeast Pacific and its larvae are distinctive. The only
other cottid larvae we have encountered with such a
heavily pigmented gut are those of our Triglops sp. which
lack ventral midline melanophores in the tail region.
that work, brief descriptions of other hemilepidotine
sculpins were also included: Hemilepidotus gilberti (as
H. g. gilberti) (7.5, 11.4, 17.5 mm), H. jordani (6.4, 10.7,
13.0 mm), and Melletes papilio (10.7, 13.7 mm) with
some comments on H. zapus (as H. g. zapus). Young of
H. hemilepidotus, ~20 mm, were described by Peden
(1964) who also described young (~20 mm) of H. jordani,
H. spinosus, and H. zapus. Young of H. gilberti (7.1,
11.6, 19.2, 24.8, 32.5 mm) were described by Hattori
(1964).
Hemilepitodus hemilepidotus (Tilesius)
(Figures 21, 22; Tables 29, 30)
Literature. Gorbunova (1964) illustrated (7.25, 10.5
mm) and briefly described larvae ofH. hemilepidotus. In
Distinguishing Features. Newly hatched, reared larvae
are ~5-6 mm long (Dunn footnote 5) and the smallest
larvae in our collections are about that size. By ~19-20
mm specimens have adult meristic complements and
have acquired a juvenile appearance, although speci-
mens as large as 23 mm have been captured pelagically
Table 29. — Meristics from young of Hemilepidotus hemilepidotus. (Specimen between dashed lines is undergoing notochord flexion.)
[* = Specimen stained with Alizarin Red S; N = Not examined.]
Anal
fin
rays
Pectoral
fin rays
Left Right
Pelvic fin
spine
and rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
Caudal fin rays
Dorsal Ventral
Vertebrae
Abdom- Cau-
inal dal
Branchi-
ostegal
rays
Dorsal
scale
rows
Scales
above
Body Dorsal fin
length Spines Rays
Second-
ary
Prin-
cipal
Prin-
cipal
Second-
ary
lateral line
Left Right
5.8 — —
5.9 — —
—
— N
— N
—
—
14
14
N
N
N
N
N
N
N
N
N
N
N
N
N
N
—
— N
— N
9.1
—
— N
-
2
9
N
N
N
N
N
N
N
-
— N
10.7
11.4*
11.5
19.0
23.0*
IX
XI
XI
XI
19
17
18
20
19
15
12
15
16
17
17
13
17
10
16
N
13
N
10
10
Buds
Buds
Buds
1,4
1,4
II
N
7
2
N
N N
6 6
N N
N N
6 6
N N
12 19
N N
N N
12 24
19
27
N
N
23
34
at the surface. Thus there may be a pelagic juvenile
phase similar to that of rockfishes, Sebastes spp. The
smallest transformed specimen collected in Oregon tide-
pools was 21 mm SL. In our collection we have no speci-
mens between ~11.5 and 19 mm.
Newly hatched larvae have pigment on the head and
along the dorsal midline of the body extending to all but
the posterior 3d or 4th myomere. An unpigmented area
occurs on the dorsal midline between about myomeres 4-
11 which fills in with pigment by ~6-7 mm. The dorso-
lateral surface of the gut is moderately pigmented. A
series of melanophores extends along the ventral midline
beginning at about the 9th or 10th postanal myomere
extending to all but the last 3 or 4 myomeres. Mela-
nophores in this series usually number <15. With devel-
opment, melanophores are added over the head, an-
terodorsal body surface, and the gut. Some internal
melanophores appear to extend dorsally from the ven-
tral midline melanophores in larvae as small as 6 mm.
Additional internal melanophores appear ventrolat-
erally between the notochord and ventral midline. By 9
mm, internal melanophores can be seen in a row above
Hemilepidotus hemilepidotus
5.8mm NL
5.9mm SL
9.1mm NL
Figure 21.— Larvae of Hemilepidotus hemilepidotus.
35
Table 30. — Measurements (mm) of young of Hemilepidotus hemilepidotus. (Specimen between dashed lines is under-
going notochord flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
5.8
5.9
0.90
1.0
0.06
0.27
0.52
0.58
2.0
2.1
1.0
0.86
0.32
0.34
—
-
9.1
2.0
0.27
0.9
3.4
1.7
0.62
0.22
0.10
10.7
11.5
19.0
2.7
3.7
6.7
0.47
0.96
1.28
1.2
1.3
2.2
4.6
6.4
11.0
2.6
3.3
6.0
1.1
1.7
4.0
0.39
0.51
0.9
0.27
BR1
0.31
BR = Broken.
Hemilepidotus hemilepidotus
I0.7mm SL
II. 5mm SL
19.0mm SL
Figure 22. — Young of Hemilepidotus hemilepidotus.
36
and below the notochord along much of the body. These
internal melanophores remain visible until the lateral
musculature obscures them. External melanophores are
not added laterally until larvae are >11 mm, and then as
a relatively light scattering compared to H. spinosus.
The ventral midline melanophores gradually become in-
ternal but remain visible through the skin and serve as a
diagnostic character to distinguish them from H. spino-
sus until dorsal scale rows develop (see Discussion). By
19 mm the entire body except the caudal peduncle and
ventral body margin are covered with pigment which is
darker dorsally and lighter below.
Four prominent spines develop on the preopercle, some
of which are visible by 9 mm. A parietal spine can be seen
at ~9 mm and later a nuchal spine, the two fusing to-
gether late in the larval period. A postocular spine
develops over the eye ~9 mm and remains obvious at
least until 11 mm, probably disappearing during trans-
formation. This spine develops only as a minute spinule
for a brief period, ~8-9.3 mm in H. spinosus, barely visi-
ble without staining. With development three spines ap-
pear in the posttemporal-supracleithral region, being
reduced to bumps in larger specimens. A cleithral spine
is apparent by ~ 19-20 mm. A nasal spine develops on the
snout. Tips of 3 spines (midopercular, interopercular,
subopercular) can be seen on the margin of the gill cover
by ~11 mm.
Relative body depth increases with development from
~15-18 to 32% SL as does snout to anus distance from
~35 to 58%. Pectoral fin length increases to ~21% SL by
19 mm. Adult complements of fin rays, dorsal scale rows,
and scales above the lateral line are countable in the 19
mm specimen and probably in smaller specimens based
on H. spinosus in which adult complements are attained
by ~14 mm. Gill membranes are noticeably free from the
isthmus in the 19 mm specimen.
Discussion. The relative position of the first postanal
ventral midline melanophore and the posterior extent of
the ventral series serve as excellent diagnostic charac-
ters to separate larvae of H. hemilepidotus from H. spi-
nosus until the dorsal scale rows are formed. In H. hemi-
lepidotus the ventral midline melanophore series begins
at about the 9th or 10th postanal myomere and consists
of usually <15 melanophores. In H. spinosus this ven-
tral row begins immediately behind the anus and con-
sists of usually >15 melanophores in all but the largest
specimens. In larger specimens, the number of dorsal
scale rows separate the two species, 4 or 5 in H. hemi-
lepidotus and 6-8 in H. spinosus. These can be counted in
specimens as small as 14 mm.
The placement and number of ventral midline mela-
nophores may not serve to distinguish H. hemilepidotus
from other species, e.g., H. gilberti, H. jordani, and H.
zapus, based on Gorbunova's (1964) and Hattori's (1964)
descriptions. All four species reportedly have the same
ventral midline melanophore pattern. And small larvae
of all four reportedly have an unpigmented break in the
dorsal midline melanophore series which fills in with
development. Additional characters given by Gorbunova
do not appear to be adequate to distinguish larvae among
these four species. One character used by Gorbunova to
distinguish late larvae of//, gilberti, i.e., presence of a
row of scales above the anal fin, is invalid. We have
observed such scales in H. spinosus and H. hemilepido-
tus, and Peden (1964) has shown them in young ofH. jor-
dani and H. zapus. These become overgrown on larger
fish. Gorbunova stated that larvae of H. gilberti develop
5 preopercular spines which would help distinguish them
but this is probably an error in observation, based on our
knowledge of other cottid larvae and illustrations by
Hattori. Pigment patterns as reported by Gorbunova
seem inadequate to distinguish species. Closer attention
to detail in exact numbers of ventral melanophores and
their placement with respect to myomere number to-
gether with a more detailed examination of external lat-
eral pigment and internal pigment along the notochord
may help resolve the problems of larval identification.
Meristic and morphometric characters as discussed by
Peden will be useful in identifying larger specimens,
> 18-20 mm. Relative body proportions may also be of
some help but these must be better defined to evaluate
their usefulness in young specimens.
Larvae (10.7, 13.7 mm) of another hemilepidotine
sculpin from the Bering sea, Melletes papilio, as de-
scribed by Gorbunova (1964) are easily distinguished by
the extension of the dorsal and ventral midline melano-
phores onto and around the urostyle.
See also Discussion under H. spinosus.
Hemilepidotus spinosus (Ayres)
(Figures 23, 24; Tables 31, 32)
Literature. Follett (1952) illustrated (12, 21 mm SL)
and briefly described several specimens of H. spinosus
which we confirm. Peden (1964) also discussed young
(~20 mm) of this species. (See also Literature section un-
der H. hemilepidotus.)
Distinguishing Features. The smallest larvae in our col-
lections are ~5 mm and appear to be recently hatched.
Specimens as large as 27 mm have been captured pelag-
ically although most juvenile characteristics appear to be
present by ~ 19-20 mm.
The smallest larvae examined, ~5 mm, have pigment
on the head and in a continuous line on the dorsal body
margin from the head to the posteriormost myomere.
Some pigment occurs on the dorsolateral surface of the
gut. A row of >15 melanophores extends along the ven-
tral body midline from immediately behind the anus to
the posteriormost myomere. A single melanophore is
usually apparent posterior to this line about midway to
the notochord tip. This melanophore later appears on the
midcaudal fin base between the upper and lower hy-
pural elements. With development external melano-
phores are added on the head, over the gut and laterally
in H. spinosus larvae as small as 6 mm in contrast to H.
hemilepidotus. By 8-9 mm, internal pigment is notice-
able in a line above the notochord along most of the
37
Hemilepidotus spinosus
6.6mm NL
8.9mm NL
Figure 23. — Larvae of Hemilepidotus spinosus.
body. Internal pigment never becomes obvious below the
notochord as in H. hemilepidotus, possibly because of
thicker musculature or because the more intense external
lateral melanophores obscure it. Pigment increases over
the dorsal body surface first anteriorly in the nape region
then posteriorly extending laterally until most of the
body is pigmented, except ventrally and on the caudal
peduncle, by ~ 19-20 mm. The ventral midline melano-
phores become embedded but remain visible through the
skin at least to ~14 mm, which serve to distinguish these
larvae from H. hemilepidotus.
Four preopercular spines develop, beginning ~7 mm.
A parietal spine can be seen ~7 mm and later a nuchal
spine develops posterior to it fusing with it in the largest
specimens. A minute postocular spine develops over the
eye in larvae 8.0-9.3 mm and then disappears. This spine
never becomes prominent as in H. hemilepidotus and is
barely visible without staining. Three spines develop in
the posttemporal-supracleithral region but become
reduced to bumps. A nasal spine and a cleithral spine are
present by 19 mm. Three spine tips (midopercular, sub-
.'58
Hemilepidotus spinosus
II. Omm SL
<■*%*,■&■**!*
1 1.8mm SL
19.0mm SL
Figure 24. — Young of Hemilepidotus spinosus.
opercular, interopercular) are visible on the gill cover
margin by ~11-12 mm.
Body depth increases from ~ 15-16% to -32-34% SL
with development as does snout to anus distance, 38-39%
to 60-61% SL. Pectoral fin length increases to ~26% SL
by 19-20 mm. Adult complements of fin rays, dorsal scale
rows, and scales above the lateral line are present by ~14
mm. In 19-20 mm specimens the gill membranes are
noticeably attached to the isthmus.
Discussion. Early life history strategies appear to differ
between H. spinosus and H. hemilepidotus. Larvae of H.
spinosus are much more abundant in our plankton sam-
ples than H. hemilepidotus and they have been collected
farther offshore, as far as 111 km from the coast. We have
taken late larvae-early juvenile stages of both species in
neuston collections but H. spinosus is much more com-
mon. Collections in Oregon tidepools have yielded pri-
marily young juveniles of H. hemilepidotus and only
rarely H. spinosus. Our data indicate that H. hemilepi-
dotus generally spawns closer to shore and the larvae are
not as neustonic as H. spinosus. Tidepools may be impor-
tant nursery areas for juvenile H. hemilepidotus whereas
39
Table 31.— Meristics from young of Hemilepidotus spinosus. (Specimen between dashed lines is undergoing notochord flexion.)
[* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; B = Bases only; N = Not examined]
Ventral
Caudal fin rays
Scales
Dorsal fin
Anal
fin
Pectoral
fin rays
Pelvic fin
spine
Preoper-
cular
midline
melano-
Dorsal
Ventral
Vertebrae
Abdom- Cau-
Branchi-
ostegal
Dorsal
scale
ab
later
sve
Body
Second-
Prin-
Prin-
Second-
il line
length
Spines
Rays
rays
Left
Right
and rays
spines
phores
ary
cipal
cipal
ary
inal
dal
rays
rows
Left
Right
5.0
24
N
N
N
N
N
N
N
_
5.7*
—
—
—
—
—
—
—
N
—
—
—
—
—
—
—
6.2
29
N
N
N
N
N
N
N
—
N
6.6
34
N
N
N
N
N
N
N
—
6.6
2
29
N
N
N
N
N
N
N
—
N
7.2
2
27
N
N
N
N
N
N
N
—
-
N
7.6*
_
—
—
—
—
—
—
N
_
_
8.0*
2
N
8.3
—
—
14B
—
—
—
3
24
N
N
N
N
N
N
N
—
N
8.6
-18B-**
15B
—
—
—
3
31
N
N
N
N
N
N
N
—
—
N
8.8*
—
—
—
6
6
—
4
N
—
4
4
—
12
18
6
—
—
8.9
-18B-**
14B
—
—
—
3
26
N
N
N
N
N
N
N
—
—
—
9.1
-19
**
15
—
—
—
4
27
N
N
X
X
N
N
N
—
—
N
9.3*
—
—
—
8
7
—
4
N
—
4
5
—
12
21
6
—
10.1
-20
**
15
1 I
X
—
4
N
N
X
N
X
N
N
-
-
N
10.4*
XI
19
15
15
15
1,1
4
4
6
6
3
12
24
6
10.6
-19
**
14
14
N
—
4
21
N
N
N
N
N
N
N
—
—
N
11.0
XI
19
15
15
15
Buds
4
22
N
N
N
X
N
N
N
—
—
—
11.0
X
20
15
15
15
Buds
4
25
N
N
N
X
N
N
N
—
—
N
11.2
XI
18
15
15
15
Buds
4
21
X
N
N
X
N
N
N
—
—
N
11.8*
XI
19
16
15
15
1,4
4
N
7
6
6
6
VI
24
6
—
—
—
11.8
XI
18
15
15
15
1,4
4
22
N
N
X
X
N
N
N
—
—
—
11.8
XI
in
14
15
15
Buds
4
18
N
N
N
X
N
N
N
—
—
N
12.0*
XI
20
15
15
15
1,4
4
N
7
6
6
3
12
24
6
—
—
12.6*
XI
19
16
16
16
1,4
4
N
7
6
6
5
12
24
6
—
—
—
13.1*
XI
19
16
15
15
1,4
4
N
8
6
6
7
L2
24
N
4
32
32
13.4
XI
19
15
15
15
Buds
4
19
N
N
N
N
N
X
N
—
—
N
13.6
XI
L9
15
15
15
1,4
4
24
N
N
N
N
N
N
N
—
—
N
13.9
XI
19
15
15
15
1,4
4
21
N
N
N
N
N
N
N
6
~30
N
14.4
XI
19
15
15
15
1,4
4
22
N
N
N
X
X
N
N
6
29
N
15.3*
XI
19
16
15
15
1,4
\
N
8
6
6
7
12
24
6
6
31
33
15.7
XI
19
15
15
15
1,4
4
13
N
N
X
X
X
X
X
6
24
N
19.0
XI
19
15
15
15
1,4
4
—
N
N
X
X
X
N
N
6
N
29
20.4*
XI
19
17
15
15
1,4
4
N
8
6
6
7
12
24
6
6
26
26
20.8
XI
20
15
15
N
1,4
4
5
X
N
N
N
N
N
X
6
21
N
22.0*
XI
18
17
16
16
1,4
4
N
8
6
6
7
12
24
6
6
34
30
juvenile H. spinosus utilize different areas, perhaps far-
ther offshore, as nurseries.
Nautichthys oculofasciatus (Girard)
(Figure 25; Tables 33, 34)
Literature. Larvae of this species were illustrated (7.5,
13 mm SL) and briefly described by Blackburn (1973)
and young stages (9.5, 13, 17, 26 mm TL) by Marliave
(1975).
Distinguishing Features. Larvae hatch at a relatively
large size, probably ~7 mm SL. The largest pelagic
specimens reported are ~16-17 mm in which all fins are
developed except the pelvics. A transformed juvenile, 26
mm TL, was taken in a benthic habitat (Marliave 1975)
indicating transformation probably occurs between these
sizes.
The larvae are distinctively pigmented over the lat-
eral surface of the body, except for the caudal peduncle,
with melanophores extending onto the dorsal and anal
finfolds and fins in the middle of the tail region. Melano-
phores also occur over the head, at the base of the pec-
toral fin, on the pectoral fin rays as a band near the fin
margin, and over the dorsolateral surface of the gut. Pig-
ment is added to the spinous dorsal fin as the anterior
rays become elongate.
Four spines develop on the preopercle appearing as
bumps. Enlarged bumps also develop in the parietal
region, anterior to it along the parietal ridge, in the
postocular region, nasal region, and several in the post-
temporal-supracleithral area. None of these appear as
pronounced sharp spines.
The length of the pelvic fins is striking, 42 and 59% SL
in our two specimens. Snout to anus length is relatively
long, 56 and 51% SL while body depth at the pectoral fin
base is relatively slender, 21 and 23% SL. The number of
myomeres (>40) is relatively high for a cottid. The larvae
have an agonidlike appearance.
Discussion. Only one species of Nautichthys occurs off
40
Table 32.
-Measurements (mm) of young of Hemilepidotus spinosus. (Specimens between dashed lines are undergoing
notochord flexion.)
Body depth
2d preoper-
Body
Head
Snout
Eye
Snout to
at pectoral
Pectoral
cular
Parietal
length
length
length
diameter
anus length
fin base
fin length
spine length
spine length
5.0
0.88
0.12
0.46
1.9
0.82
0.24
—
—
6.2
1.1
0.16
0.56
2.6
0.94
0.31
—
—
6.6
1.0
0.24
0.54
2.5
0.98
0.35
—
—
6.6
1.4
0.24
0.56
2.6
1.0
0.47
—
0.04
7.2
1.4
0.31
0.62
2.8
1.2
0.47
—
0.04
8.3
1.9
0.39
0.82
3.6
1.6
0.47
0.24
0.10
8.6
1.6
0.35
0.78
3.5
1.5
0.66
0.22
0.12
8.9
1.7
0.23
0.82
3.5
1.7
0.74
0.27
0.20
9.1
1.9
0.39
0.90
3.8
1.8
0.78
0.27
0.20
10.1
2.3
0.40
1.2
3.9
2.0
0.82
0.39
0.23
10.6
2.5
0.39
1.2
4.5
2.1
0.82
0.35
0.31
11.0
3.1
0.39
1.3
5.4
2.7
1.5
0.58
0.23
11.0
3.1
0.48
1.3
5.5
3.0
1.5
0.70
-0.27
11.2
3.3
0.40
1.6
5.5
3.0
1.6
0.66
0.31
11.8
3.8
0.70
1.4
6.2
3.4
1.8
0.72
0.24
11.8
3.8
0.80
1.6
6.7
3.4
1.9
0.78
0.31
13.4
4.7
0.96
1.9
7.0
4.2
2.4
0.88
-0.2
13.6
4.6
0.96
2.0
7.4
4.2
2.7
0.88
—
13.9
5.1
1.0
1.8
7.4
4.3
2.9
0.9
—
14.4
5.4
1.0
1.8
8.2
5.0
2.9
BR1
—
15.7
5.7
1.4
2.0
9.3
5.0
3.4
1.1
—
19.0
7.4
1.7
2.2
11.5
6.4
4.8
1.1
—
20.8
7.8
2.2
2.2
12.5
6.6
5.4
1.1
—
'BR = Broken.
Nautichthys oculofasciatus
II. 7mm NL
Figure 25. — Larvae of Nautichthys oculofasciatus.
41
Oregon and the larvae are striking in appearance. The
larvae are rare in our collections. Only two specimens
have been taken, both 9 km off Newport, Oreg., in
March. Larvae of two additional northeast species N.
pribilovius and N. robustus are unknown.
Table 33. — Meristics from larvae of Nautichthys oculofasciatus.
(Specimen above dashed line is undergoing notochord flexion.)
Dorsal Dorsal Anal Pectoral
Ventral
Pelvic fin Preoper- midline
Body
length
fin
spines
fin
rays
fin
rays
fin rays
Left Right
spine
and rays
cular
spines
melano-
phores
11.7
vn
28
19
13 12
-
-
N2
16.6
IX
28
19
14 N
—
4
N
'Not present as a distinct row.
2N = Not examined.
Radulinus asprellus Gilbert
(Figures 26, 27; Tables 35, 36)
Literature. Larvae of this species have not been pre-
viously described.
Distinguishing Features. The smallest larvae we have
collected are ~4.5-5 mm long and probably recently
hatched. The largest specimens captured pelagically,
~ 14-15 mm, have developed most juvenile character-
istics including adult fin complements, increased
pigmentation, and lateral line scales.
Larvae of R. asprellus are one of the most heavily
pigmented among cottids. Melanophores occur over most
Table 34. — Measurements (mm) of larvae of Nautichthys oculofasciatus. (Specimen above dashed line is undergoing
notochord flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper -
cular
spine length
Parietal
spine length
11.7
3.4
0.80
1.1
6.6
2.5
4.9
-
-
16.6
4.5
1.1
1.3
8.4
3.8
9.8
0.20
—
Table 35. — Meristics from young of Radulinus asprellus. (Specimens between dashed lines are undergoing notochord flexion.)
[* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; *** = Present but not countable as
distinct melanophores; B = Bases only; D = Damaged; N = Not examined.]
Pelvic Ventral Caudal fin rays
Anal Pectoral fin Preoper- midline*** Dorsal Ventral Vertebrae Branchi-
Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal
length Spines Rays rays Left Right &rays spines phores ary cipal cipal ary inal dal rays
4.7 — — — — — NNNNNNNN
5.2 — — — — — — N NNNN NNN
5.9* _____ N N
6.1 NNNNNNNN
6.7 — — — — — — — N NNNN NNN
7.2 — — — — — — 3 N NNNN NNN
7.8* _____ _ N N ____ ___
7.9 — — — — — — — N NNNN NNN
8.5* _____ N N — - — ___
9.1 -22B-** 18B — — NNNNNNNN
9.6 VII 21 22 — — 3 N NNNN NNN
10.1 -21-** 22 18N NNNNNNNN
10.9 X 22 22 19 N Buds 4 N NNNN NNN
11.4
-23-
**
22
18
18
Buds
4
N
N
N
N
N
N
N
N
11.4*
X
21
23
—
—
—
N
N
—
5
5
—
—
—
—
11.5
IX
22
24
18
18
Buds
4
N
X
N
N
N
N
N
N
11.7*
X
22
22
16
N
—
N
N
—
5
5
—
12
20
6
12.0
IX
22
23
17
17
Buds
4
N
N
N
N
N
N
N
N
12.0*
X
21
24
18
19
1,3
N
N
—
6
6
—
12
23
6
12.6
IX
20
22
19
18
1,3
4
X
N
X
X
N
N
N
N
12.6
X
23
24
19
19
1,3
4
N
X
N
N
N
N
N
N
13.0
IX
22
24
18
19
Buds
4
N
N
N
N
X
N
N
N
13.0*
vm
D
23
18
18
1,3
N
N
—
6
6
—
12
27
6
13.9*
X
22
23
18
18
1,3
N
X
3
6
6
2
12
27
6
14.1
IX
23
23
18
L8
1,3
4
N
X
N
N
N
N
N
N
14.1
X
22
23
19
19
1,3
4
N
N
X
N
N
N
N
N
14.4
IX
22
24
18
18
1,3
4
N
N
N
N
N
N
N
N
14.4
X
23
24
19
19
1,3
4
N
N
X
N
N
N
N
N
14.7*
IX
22
23
19
19
1,3
N
N
3
6
6
2
12
27
6
15.2*
IX
23
24
18
18
1,3
N
N
3
6
6
2
12
27
6
42
Radulinus asprellus
4.7mm NL
7.9mm NL
■ Mas V«+>*' 2**£ s#
<*&>
9.6mm NL
Figure 26. — Larvae of Radulinus asprellus.
of the body including the head, most of the gut, along
dorsal and ventral midlines, and laterally except dorso-
lateral^ above the body midline over the gut region and
dorsally and laterally on the tail tip. This pigment is
somewhat darker along the dorsal and ventral body
midlines. Some ventral midline melanophores extend
beyond the end of the lateral body pigment onto the tail
tip, particularly in smaller specimens. A distinctive
series of elongate melanophores occurs along the lateral
midline appearing almost as a solid line. The presence of
these melanophores help distinguish larvae of R. asprel-
lus from Scorpaenichthys marmoratus which they resem-
ble at small sizes. Pigmentation generally increases with
development with only the tail tip or caudal peduncle re-
maining unpigmented. The lateral midline pigment is
noticeable on all but the largest specimens.
Four small spines develop along the preopercular
margin, visible by ~ 10-11 mm. They never become
prominent. A parietal and nuchal spine develop as 2
minute spines which soon appear only as bumps. A nasal
spine is evident on the largest transforming specimens
~14-15 mm.
The gut is rather distinctively coiled with snout to
anus length ranging from ~44 to 54% SL. The snout
becomes quite pointed further helping to distinguish
these larvae from S. marmoratus. The larvae are rela-
tively slender with body depth at the pectoral fin base
never exceeding 25% SL during the pelagic phase and
always <20% SL in specimens <10 mm. The adult com-
plement of all fin rays or ray elements is attained by ~11-
12 mm.
Discussion. Three species of Radulinus occur in the
northeast Pacific. Larvae of R. boleoides are described in
43
Radulinus asprellus
,.y
I0.9mm NL
12.6mm SL
14.4mm SL
Figure 27. — Young of Radulinus asprellus.
the next section. They are distinguishable from R.
asprellus on the basis of body proportions and small pig-
ment differences as given in that section, at least for the
size range described. Larvae of the third species, R.
vinculus, are unknown.
Larvae of R. asprellus are one of the more heavily
pigmented cottids along with, e.g., Scorpaenichyths
marmoratus, Rhamphocottus richardsoni, and Blepsias
cirrhosus (Blackburn 1973; Marliave 1975), all of which
have melanophores covering most of the body except the
tail tip or caudal peduncle. Interestingly only one of
these, S. marmoratus, is known to be common in the
neuston, indicating larval pigmentation may not reflect
habitat preference, at least in these species.
Radulinus boleoides Gilbert
(Figure 28; Tables 37, 38)
Literature. The larvae of this species have not been
previously described.
Distinguishing Features. We have identified only two
larvae as R. boleoides in our collections, 7.5 and 8.7 mm
long. At comparative sizes they are somewhat more ad-
vanced in development than larvae of R. asprellus.
The pigment pattern closely resembles that of R.
asprellus with melanophores covering most of the body
laterally except the tail tip. A few ventral midline mela-
44
Table 36. — Measurements (mm) of young of Radulinus asprellus. (Specimens between dashed lines are undergoing
notochord flexion.)
Body depth
2d preoper-
Body
Head
Snout
Eye
Snout to
at pectoral
Pectoral
cular
Parietal
length
length
length
diameter
anus length
fin base
fin length
spine length
spine length
4.7
1.1
0.10
0.51
2.6
0.86
0.47
—
—
5.2
1.2
0.20
0.47
2.4
0.82
0.27
—
—
6.1
1.3
0.32
0.54
2.8
0.98
0.27
—
—
6.7
1.6
0.35
0.62
3.3
1.1
0.35
—
—
7.2
1.9
0.43
0.58
3.4
1.2
0.35
—
—
7.9
1.8
0.39
0.70
3.5
1.3
0.55
—
—
9.1
2.3
0.56
0.56
4.2
1.6
0.55
—
—
9.6
2.7
0.82
0.82
4.6
1.8
0.78
—
—
10.1
2.4
-0.50
0.88
4.9
1.8
0.80
—
—
10.9
2.9
0.78
1.0
5.9
2.6
1.1
—
—
11.4
3.0
0.72
1.0
6.0
2.6
1.8
—
—
11.5
3.5
0.88
1.0
6.2
2.5
2.0
0.27
—
12.0
3.6
1.0
1.2
6.3
2.5
2.1
0.23
—
12.6
4.1
0.88
1.4
6.6
3.0
3.1
0.35
—
12.6
3.7
0.88
1.21
6.6
3.0
l.H
0.23
—
13.0
3.6
0.96
1.1
6.2
2.6
1.7
0.27
—
14.1
4.5
0.96
1.4
7.6
3.5
2.0
0.40
—
14.1
4.8
1.3
1.4
7.0
2.8
2.7
0.46
—
14.4
4.7
1.1
1.5
7.0
3.0
3.7
0.43
—
14.4
4.2
1.0
1.4
7.0
3.4
2.2
0.35
—
Radulinus boleoides
'jfJjA.
&C-SJ
8.7mm NL
Figure 28. — Larva of Radulinus boleoides.
Table 37.
-Meristics from larvae of Radulinus boleoides.
mens are undergoing notochord flexion.)
(Speci-
Ventral
Dorsal
Dorsal Anal
Pectoral
Pelvic fin
Preoper-
midline
Body
fin
fin fin
fin rays
spine
cular
melano-
length
spines
rays rays
Left Right
and rays
spines
phores
7.5
VIII
20 15
18 N2
Buds
—
N
8.7
IX
16 20
19 N
1,3
—
N
'Present but not countable as distinct melanophores.
2N = Not examined.
nophores extend onto the tail tip. They differ somewhat
in the posterior extent of the lateral pigment which is less
in R. boleoides than in R. asprellus of similar size, i.e., a
greater portion of the tail tip is pigmentless in R.
boleoides. Larvae of R. boleoides also have a distinctive
series of melanophores along the lateral midline as do R.
asprellus.
No head spines are visible on the two flexion larvae in
our collection, although they may develop later.
Table 38. — Measurements
(mm) of larvae of Radulinus boleoides.
(Specimens are
undergoing notochord flexion.)
Body Head
length length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
7.5 2.4
8.7 2.6
0.58
0.64
0.70
0.78
4.4
5.0
1.7
2.0
0.98
1.2
—
—
45
The coiled gut is rather prominent and distinctive, as
in R. asprellus, but snout to anus length is considerably
greater in R. boleoides (58% SL) than R. asprellus (44-
47% SL) of similar size. Body depth at the pectoral fin
base is also greater in R. boleoides (22-23% SL) than R.
asprellus (16-18% SL). The pectoral fin is relatively
longer in R. boleoides (13-14% SL) compared to R.
asprellus at that size (5-7% SL). Both specimens of R.
boleoides are undergoing notochord flexion and adult fin
ray complements have not been attained, explaining the
discrepancies in the fin ray counts in Table 37.
Discussion. Identification was based on the strong sim-
ilarity of these larvae to R. asprellus and the major dif-
ferences in body proportions which were obviously not
caused by distortion. The only other species of Radu-
linus which occurs off Oregon is R. boleoides and it is
relatively rare, explaining the paucity of larvae in our
collections. Our two specimens were taken 9 and 18 km
off the Oregon coast in March and April.
Rhamphocottus richardsoni Gunther
(Figure 29; Tables 39, 40)
Literature. Larvae (6.7, 10 mm SL) were illustrated
and briefly described by Blackburn (1973) and young
(10, 11.5, 15 mm TL) by Marliave (1975).
Distinguishing Features. Hatching occurs ~6-7 mm
and transformation at ~ 14-15 mm (Blackburn 1973;
Marliave 1975). Pelagic specimens in our collections
range from 6.0 to 11.8 mm.
Larvae are rather heavily pigmented with melano-
phores covering most of the body except the tail tip or
caudal peduncle. Pigment covers the cheeks, and ventral
surface of the gut on the smallest specimens. Pigment
does not appear noticeably darker along the dorsal and
ventral body margins and no distinct line of melano-
phores occurs along the lateral midline. Some melano-
phores occur on the ventral finfold and occasionally on
the dorsal finfold. The preanal finfold is darkly
pigmented. Melanophores are present on the base of the
pectoral fin. The caudal peduncle remains unpigmented
through the pelagic period. Body pigment changes little
with development except as transformation approaches,
pigment is added to the spinous dorsal fin and pecto-
ral fin base, and pigment bands begin to show on the
body.
Only 1 spine develops on the dorsal margin of pre-
opercle, evident by ~10 mm. Smaller spiny projections
appear along the preopercular margin which are modi-
fied scales developing over the head region. A postocular
spine is apparent over the eye by ~8 mm and becomes
rather pronounced in larger specimens. A parietal and a
nuchal spine develop and fuse together appearing as one.
Two spines become obvious in the posttemporal-supra-
cleithral region and a cleithral spine and nasal spine are
evident on our largest specimens.
Body shape is characteristic. Larvae are deep bodied,
increasing from 29 to 40% SL between 8 and 12 mm.
Snout to anus distance is >60% SL. By 10 mm, larvae
begin to resemble adults in form. The larvae have a dis-
tinct pigmented preanal finfold which remains evident
anterior to the anal opening at least until ~12 mm. To
our knowledge, a preanal finfold has been reported in
Scorpaenichthys marmoratus (O'Connell 1953), Hemi-
tripterus americanus (Warfel and Merriman 1944; Fui-
man 1976), possibly H. villosus as indicated on a figure
by Okiyama and Sando (1976), Myoxocephalus quadri-
cornis, M. scorpius (Khan 1972), possibly M. aenaeus
based on figures by Lund and Marcy (1975), possibly
Gymnocanthus tricuspis and G. herzensteini based on
illustrations by Rass (1949) and Kyushin (1970), Eno-
Table 39.— Meristics from larvae of Rhamphocottus richardsoni. (Specimen above dashed line is undergoing notochord
flexion.)
(* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; *** = Present but not countable as
distinct melanophores; B = Bases only; N = Not examined.]
Dorsal fin
Spines Rays
Anal
fin
rays
Pectoral
fin rays
Left Right
Pelvic
fin
spine
& rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
Caudal fin rays
Vertet
Abdom-
inal
rae
Cau-
dal
>** Dorsal
Second- Prin-
ary cipal
Ventral
Branchi-
Body
length
Prin-
cipal
Second-
ary
ostegal
rays
8.4
-14B-**
6B
-
-
Buds
-
N
\
N
N
N
N
N
N
10.6
11.7
11.8*
VIII 13
VIII 12
VIII 13
6
6
7
16
15
16
16
16
16
1,3
1,3
1,3
1
1
1
N
N
N
N
N
1
N
N
6
N
N
5
N
N
1
N
N
12
N
N
15
N
N
6
Table 40. — Measurements (mm) of larvae of R hamphocottus richardsoni.
notochord flexion.)
(Specimen above dashed line is undergoing
Body
length
Head
length
2.6
Snout
length
0.55
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
8.4
0.94
5.5
2.4
1.0
-
—
10.6
11.7
4.1
4.8
1.0
1.0
1.1
1.3
7.7
8.1
4.0
4.7
2.2
2.6
—
0.43
0.35
46
Rhamphocottus richardsoni
£2zr~/ r™
•>3«^*W
3V»*'lS.*irV3*>.~
\" ' /'k-~- 1 ** * *'■*' • ' <; @
8.4mm NL
mj$$™-:-
10.6mm SL
II. 7mm SL
Figure 29. — Young of Rhamphocottus richardsoni.
47
phrys bison (Misitano 1978), and E. bubalis and possibly
E. lilljeborgi (Russell 1976), within the Cottidae.
Discussion. Rhamphocottus is a monotypic genus. The
larvae are distinctive. They share the character of having
a rather heavily pigmented body, except for the tail tip,
with a number of other northeast Pacific species, as dis-
cussed under Radulinus asprellus, but their character-
istic body shape renders them outstanding except
perhaps at the smallest sizes. The heavily pigmented
preanal finfold, and presence of pigment on the anal fin-
fold, base of the pectoral fin, and the ventral surface of
the gut help distinguish small larvae. The body becomes
covered with distinctive spinelike scales while still in the
pelagic habitat. The larvae are relatively rare in our col-
lections with only seven identified specimens, all taken
2-9 km off the Oregon coast in March and April.
Scorpaenichthys marmoratus (Ayres)
(Figures 30, 31; Tables 41, 42)
Literature. Eggs and young (5.85, 6.26, 10, 17, 30, 48
mm) of this species were described by O'Connell (1953).
Distinguishing Features. Larvae are between 4 and 6
mm long at hatching. Most of our pelagic specimens are
<15 mm. By ~13-14 mm they have a juvenile appear-
ance and fully formed fins. However, we have captured a
21.6 mm specimen pelagically and a 35 mm specimen in
a neuston tow 9 km off Newport, Oreg. O'Connell (1953)
reported specimens as large as 29 mm were collected with
dip nets.
Larvae are very heavily pigmented over most of the
body, usually appearing darker than larvae of Radulinus
Scorpaenichthys marmoratus
5.3mm NL
7.5mm NL
8.6mm NL
Figure 30. — Larvae of Scorpaenichthys marmoratus.
48
Scorpaenichthys marmoratus
10. 4mm SL
13.8mm SL
Figure 31. — Young of Scorpaenichthys marmoratus.
asprellus or Rhamphocottus richardsoni. In small larvae
melanophores cover the head, the gut except the ventral
margin, the dorsal body margin, the ventral margin of
the tail, and the lateral body surface except for the tail
tip. The lateral pigment extends more posteriorly than in
Radulinus spp. The dorsal body margin, dorsal surface of
the gut, and postanal ventral body margin appear darker
than the rest of the body. The pectoral fin, preanal fin-
fold, and body finfold are unpigmented. With develop-
ment, melanophores are added on the head and laterally
except along the ventral margin of the gut which remains
unpigmented until ~8-9 mm and the caudal peduncle
which retains an unpigmented band at least until 14-15
mm. As the median fins develop the dorsal and ventral
body margins do not appear as heavily outlined. On
specimens ~13-14 mm a discrete melanophore can be
seen at the base of most dorsal fin soft rays. Larvae gen-
erally do not develop a distinct line of pigment along the
lateral midline as observed in larvae of Radulinus spp.
The margin of the preopercle develops 4 spines, not 3
as reported by O'Connell (1953), but only the upper 3
spines become pronounced. They are first noticeable ~8
49
mm, and become striking in postflexion stages with the
length of the 2d spine ranging from 22 to 26% HL. Raised
bumps develop in the parietal region, the posttemporal-
supracleithral region, and the cleithral region, but spines
never form. A nasal spine becomes obvious by ~ 13-14
mm.
Preflexion larvae are relatively slender with body
depth at pectoral fin base ranging from 17 to 19% SL.
Table 41. — Meristics from young of Scorpaenichthys marmoratus. (Specimens between dashed lines are undergoing notochord
flexion.)
[* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; *** = Present but not countable as
distinct melanophores; B = bases only; N = Not examined.]
Pelvic
Ventral
Caudal fin rays
Dorsal fin
Spines Rays
Anal
fin
rays
Pectoral
fin rays
fin
spine
& rays
Preoper-
cular
spines
midline*
melano-
phores
** Dorsal
Ventral
Vertebrae
Abdom- Cau-
inal dal
Branchi-
Body
Second-
ary
Prin-
cipal
Prin-
cipal
Second-
ary
ostegal
length
Left
Right
rays
5.1*
N
N
—
—
—
—
—
—
—
5.3
X
N
N
N
N
N
N
N
5.4
X
N
N
N
N
N
N
N
5.8
N
N
N
N
N
N
N
N
6.1
N
N
N
N
N
N
N
N
6.3
N
N
N
N
N
N
N
N
6.6*
N
N
7.5
N
N
N
N
N
N
N
N
7.5*
N
X
7.7*
N
N
8.1
2
N
N
N
N
N
N
N
N
8.1
-15B-**
12B
—
—
—
4
N
N
N
N
N
N
N
N
8.2*
N
N
8.6
-16B-**
13B
—
—
—
2
N
N
N
N
X
N
N
N
8.7
-15-
**
13
11
14
—
4
N
N
N
N
N
N
N
N
9.1*
12
10
N
N
—
6
6
—
14
16
6
9.3
-16-
**
13
14
15
Buds
4
N
N
N
N
X
N
N
N
10.1*
X
18
14
13
13
—
N
N
5
6
6
2
14
19
6
10.4
XI
17
12
15
15
Buds
4
N
N
N
N
N
N
N
N
10.4
XI
15
11
14
14
Buds
4
N
N
N
X
X
X
N
N
10.7
XI
17
12
15
15
Buds
4
N
N
N
N
N
X
N
N
10.9*
XI
17
14
15
15
1,3
N
N
7
6
6
6
15
21
6
11.5
X
16
12
14
15
1,5
4
N
N
X
X
X
N
N
N
12.2*
XI
17
14
16
16
1,5
N
N
9
6
6
5
15
21
6
13.8
XI
17
12
15
15
1,5
4
N
N
X
N
X
X
N
N
13.8*
XI
17
13
N
15
1,5
N
N
9
6
6
9
15
21
6
14.7
XI
15
1.3
15
15
1,5
4
X
N
N
N
X
N
N
N
21.6
X
16
13
15
15
1,5
4
N
X
N
N
N
N
N
N
Table 42. — Measurements (mm;
Body
length
of young of Scorpaenichthys marmoratus.
undergoing notochord flexion.)
(Specimens between dashed lines are
Body depth 2d preoper-
Head Snout Eye Snout to at pectoral Pectoral cular Parietal
length length diameter anus length fin base fin length spine length spine length
5.3
1.1
0.27
0.44
2.8
0.90
0.44
—
—
5.4
1.2
0.27
0.56
2.9
0.98
0.40
—
—
5.8
1.3
0.26
0.56
3.1
0.98
0.43
—
—
6.1
1.4
0.30
0.56
3.1
1.1
0.48
—
—
6.3
1.3
0.36
0.56
3.1
1.2
0.46
—
—
7.5
1.6
0.27
0.66
3.7
1.2
0.82
—
—
8.1
1.9
0.31
0.82
4.6
1.9
0.55
—
—
8.1
2.5
0.39
1.05
4.6
2.1
0.82
0.32
—
8.6
1.9
0.35
0.68
4.2
1.6
0.70
—
—
8.7
2.2
0.39
0.98
5.2
2.4
0.96
0.42
—
9.3
3.1
0.40
1.4
5.8
3.2
1.6
0.70
—
10.4
3.8
0.72
2.2
6.2
3.3
1.8
0.90
—
10.4
3.1
0.48
1.4
5.9
3.3
1.8
0.78
—
10.7
3.4
0.48
1.7
6.5
3.8
2.0
0.78
—
11.5
3.8
0.40
1.8
7.4
3.8
2.2
1.01
Bump
13.8
5.4
0.96
2.2
8.9
4.6
3.0
1.24
Bump
14.7
5.4
0.88
2.5
9.7
5.2
3.4
1.36
—
21.6
8.8
1.92
2.7
15.5
8.2
6.2
1.44
—
50
They become much deeper bodied with development
with a body depth of 32-35% SL in postflexion stages.
Snout to anus length is relatively long, increasing from
49-53% SL in preflexion larvae to 57-66% SL in post-
flexion larvae. The snout develops a rounded appear-
ance, never pointed as in Radulinus sp. An unpigmented
preanal finfold remains evident up to ~13-14 mm. The
presence of such a preanal finfold has been reported for
relatively few cottid genera (see Discussion under Rham-
phocottus richardsoni).
Discussion. Scorpaenichthys is a monotypic genus. The
larvae are rather distinctive and one of the most heavily
pigmented forms of northeast Pacific cottid species.
Small larvae appear somewhat similar to Radulinus spp.
but have lateral pigment extending further toward the
tail tip, no ventral midline melanophores posterior to the
lateral pigment and usually no distinct line of melano-
phores along the lateral midline. Occasionally newly
hatched larvae will have very few melanophores on the
lateral body surface causing them to resemble small
Hemilepidotus spinosus. However, their gut is longer and
the dorsal and ventral midline pigment does not extend
as far onto the tail tip as in H. spinosus.
Larvae of S. marmoratus appear to be strongly asso-
ciated with the neuston. O'Connell (1953) reported dip
net collections contained specimens 7.4-29 mm long, and
we have taken specimens 4.0-35 mm in neuston tows off
Oregon. One 10-min neuston tow taken 9 km off New-
port, Oreg., in March contained 829 larvae, mostly 4-5
mm long and recently hatched. Richardson and Pearcy
(1977) reported them to be a coastal form, as the larvae
were only taken within 28 km of the coast in oblique
plankton tows. However, we have since collected them in
the neuston as far as 65 km offshore, well beyond the
continental shelf-slope break.
Cottoid Type A
(Figure 32; Tables 43, 44)
Literature. No published descriptions of this form exist
although illustrations were given by Richardson and
Bond9 as "Globby Type A." These larvae were called
'Richardson, S. L., and C. E. Bond. 1978. Two unusual cottoid
fishes from the northeast Pacific. Unpubl. manuscr., 6 p. Gulf Coast
Research Laboratory, Ocean Springs, MS 39564, or Department of Fish-
eries and Wildlife, Oregon State University, Corvallis, OR 97331.
Psychrolutes-like sp. 1 by Richardson (footnote 4) and
Richardson and Pearcy (1977).
Distinguishing Features. Three larval specimens (9.8,
12.8, 13.4 mm) of this unusual looking form have been
collected off Oregon. The smallest specimen is under-
going notochord flexion and the two larger specimens are
postflexion larvae. The largest larva, which was stained,
has the full complement of dorsal, anal, pectoral, and
pelvic fin rays but secondary caudal rays are still
developing and the vertebral centra are not completely
ossified, thus transformation occurs at a larger size.
The larvae are pigmented over the entire body except
at the tail tip. The basal portion of the pectoral fin is
pigmented, but its distal portion and the dorsal and anal
finfolds are unpigmented.
No spines are apparent in the head region. The belly is
covered with small, thumb-tack prickles which extend
dorsolateral^ in lesser numbers and a few are on the tail
region in the larger specimens.
These larvae are globose in shape, unlike any other
known cottid larvae. The outer skin is loose. They are
deep bodied with depth at the pectoral fin base 42 and
56% SL in the two specimens illustrated. Snout to anus
distance is also great, 58 and 79% SL, respectively. The
pelvic fins appear to be inserted in pockets in the skin
with only the tips exposed in all three larvae. This condi-
tion may be variable as reported in adults in a Southern
Hemisphere cottoid genus Neophrynichthys (Nelson 197-
7) and as observed in larvae of a Bering Sea cottoid form
(unpubl. data) referred to as Malacocottus zonurus -
Type 1 by Richardson and Bond (footnote 9). The first
dorsal fin appears to be embedded in the skin and the
second dorsal and anal fins are partially skin covered.
Discussion. Meristics, particularly the high number of
pectoral fin rays and the possession of seven branchi-
ostegal rays, agree with only one species described from
the area, Psychrolutes phrictus (Stein and Bond 1978).
Young stages of P. phrictus <50 mm also have prickles in
the belly region similar to these larvae. Based on this
information alone our Cottoid Type A larvae would ap-
pear to be P. phrictus. However, discovery of a single 26.4
mm juvenile (Richardson and Bond footnote 9) which
also agrees with P. phrictus in meristics and possession of
belly prickles has lead to uncertainty regarding its iden-
tity. This unidentified juvenile differs drastically in body
form from a 35 mm P. phrictus, primarily in relative
Table 43. — Meristics from larvae of Cottoid Type A. (Specimen above dashed line is undergoing notochord flexion.)
= Specimen stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; D = Damaged; N = Not examined.]
Dorsal fin
Spines Rays
Anal
fin
rays
Pectoral
fin rays
Left Right
Pelvic
fin
spine
& rays
Preoper-
cular
spines
Ventral
midline
melano-
phores
Caudal
Dorsal
Second- Prin-
ary cipal
fin rays
Ventral
Vertebrae
Abdom- Cau-
inal dal
Branchi-
Body
length
Prin-
cipal
Second-
ary
ostegal
rays
9.8
-15-**
11
23
23
-
-
-
N
N
N
N
N
N
N
12.8
13.4*
-16-**
VIII 20
11
14
24
26
D
26
1,3
-
-
N
N
6
N
7
N
4
N
12
N
23
N
7
51
Cottoid Type A
9.8 mm SL
12.8 mm SL
Figure 32. — Larvae of Cottoid Type A. Arrows indicate the position of the anus.
Table 44. — Measurements (mm) of larvae of Cottoid Type A. (Specimen above dashed line is undergoing notoehord
flexion.)
Body
length
Head
length
Snout
length
Eye
diameter
Snout to
anus length
Body depth
at pectoral
fin base
Pectoral
fin length
2d preoper-
cular
spine length
Parietal
spine length
9.8
3.6
0.66
1.2
5.7
4.1
1.7
-
-
L2.8
6.1
1.5
1.7
10.1
7.2
3.7
—
—
52
body depth (51% and 31% SL, respectively) and snout to
anus length (61% and 51% SL, respectively). The
unidentified 26.4 mm juvenile agrees much more closely
with our Cottoid Type A larvae than with the 35 mm P.
phrictus juvenile. It seems unlikely that a drastic change
in body proportion would occur during the juvenile
period between 26 and 35 mm. And the 26.4 mm speci-
men does not appear to be distorted, based on radio-
graphs. It also seems unlikely that two such unusual
forms would exist in the northeast Pacific that agree so
well in meristics and belly prickles. However, the identi-
fication of our Cottoid Type A larvae as P. phrictus can-
not be positively made until additional material becomes
available to provide a conclusive linkage between our lar-
vae and identified juveniles.
If our Cottoid Type A larvae are found to be P.
phrictus, then placement of this species in the genus Psy-
chrolutes is obviously in error based on the extreme dis-
similarity of these larvae and those of P. paradoxus de-
scribed by Blackburn (1973) (10.3 mm SL) and Marliave
(1975) (10.5, 13, 14 mm TL), and its generic position
should be reexamined. Alternatively, these larvae could
represent a new species and possibly a new genus of cot-
toid fish from the northeast Pacific. The term "cottoid"
has been used here because of the dissimilarity of these
larvae to any known larvae in the Family Cottidae.
NOTES ON OTHER
NORTHEAST PACIFIC COTTID LARVAE
Larvae are known for 14 additional species of north-
east Pacific cottids not described in this paper (Table
45). Those in the genera Artedius, Clinocottus, and
Hemilepidotus (including Melletes from the Bering Sea)
were discussed under the descriptions of related species.
Larvae of Oligocottus spp. were discussed under Cottidae
Type 3. Comments on the larval characters in remaining
genera, Blepsias, Dasycottus, Gilbertidia, ?Malaco-
cottus, Myoxocephalus, Orthonopias, and Psychrolutes
are included here for comparative purposes, based on the
literature cited in Table 45 and personal observations of
specimens not in the Oregon State University Larval
Fish Reference Collection.
Larvae of Blepsias cirrhosus are one of the dark,
heavily pigmented forms along with Radulinus spp.,
Rhamphocottus richardsoni, and Scorpaenichthys
marmoratus. The entire body is covered with dense
melanophores except for the tail tip, the ventral surface
of the gut, and the cheek. The pectoral fins are un-
pigmented and no melanophores occur on the finfold as
in R. richardsoni. They have no preanal finfold as in R.
richardsoni and S. marmoratus. They apparently hatch
at a relatively large size. The smallest larvae reported
from plankton collections are 10 mm TL and just be-
ginning to undergo notochord flexion. Thus, they are
generally less developed than the above-mentioned dark
cottid larvae at comparable sizes. The larvae appear
more slender with respect to body length than S. marmo-
ratus and R. richardsoni and relative snout to anus
length is less than in R. richardsoni. They do not develop
the pronounced preopercular spines seen in S. mar-
moratus. The largest specimen reported from plankton
collections is 25.5 mm TL and is apparently trans-
formed, or nearly so.
Table 45.— Additional northeast Pacific cottid species for which larvae-and young are known.
Species
Artedius lateralis
Blepsias cirrhosus
Clinocottus analis
Clinocottus recalvus
Dasycottus setiger
Gilbertidia sigalutes
Hem ilepidotus jordani
Hemilepidotus zapus
?Malacocottus zonurus
- Type 1
Myoxocephalus polya-
canthocephalus
Oligocottus maculosus
Oligocottus snyderi
Orthonopias triads
Psychrolutes paradoxus
Reference
Budd 1940
Marliave 1975
Blackburn 1973
Marliave 1975
Eigenmann 1892
Budd 1940
Morris 1951
Blackburn 1973
Blackburn 1973
Marliave 1975
Gorbunova 1964
Peden 1964
Peden 1964
Richardson and Bond 1978
Richardson (unpubl. data)
Blackburn 1973
Stein 1972
Stein 1973
Stein 1972
Bolin 1941
Blackburn 1973
Marliave 1975
Size illustrated
4.1mmSL
4, 8, 11, 14 mm TL
12.2 mm SL
10, 14, 19, 25.5 mm TL
~4 mm
~4 mm
4.6,5.0,7.6,8.3,9.9, 10.8,18.0,
24.3 mm TL
7.4 mm SL
7.3, 9.5 mm SL
7, 13,15, 25, 34 mm TL
6.4, 10.7, 13.0 mm
~20mm
~20mm
7.0,9.8, 14.2, 24.0 mm SL
6.6, 7.0, 8.8, 9.8, 10.4, 14.2, 24.0
mm SL
7.7, 10.7 mm SL
4.6-5.2,6.0, 6.6, 9.2 mm TL
4.6-5.2, 6.0, 6.6, 9.2, 12.2 mm TL
4.5-4.75, 5.5 mm TL
~3-4 mm SL
10.3 mm SL
10.5, 13, 14, 13 mm TL
See text footnote 9.
53
Larvae of Dasycottus setiger are characterized by a
relative lack of pigment on the body except on the head
and over the relatively short gut. Melanophores in these
two regions are low in numbers but are large in appear-
ance, thus covering most of the head and abdominal
cavity. Pigment is absent from the postanal ventral
midline. Pigment is present on the underside of the pec-
toral fin and later is added to the blade and outer side of
the fin. The general pigment pattern, except for the
pigmented pectorals, resembles our Triglops sp. but the
number of myomeres is much lower for D. setiger, ~34-
35. The smallest larvae reported are 7.4 mm and pre-
sumed to be recently hatched. Fin rays are formed by 12
mm and the pigment pattern remains unchanged at least
until 19 mm. A 24 mm SL specimen observed by us has
begun to acquire juvenile pigmentation.
Small larvae of Gilbertidia sigalutes and Psychrolutes
paradoxus are similar to each other. Both hatch at ~6-7
mm. The smallest larvae resemble those of liparids in
general shape. Both have moderate melanistic pigment
over the head, nape, gut, and on the pectoral fins. Post-
anal ventral midline melanophores are absent. On both,
pigment is added to the lateral body surface above the
abdominal cavity and posterior to it, with relatively more
pigment present in P. paradoxus than G. sigalutes of
similar size particularly in larger specimens. Both
develop the appearance of having loose skin. The number
of pectoral fin rays separate the two with > 19 for P. para-
doxus and <18 for G. sigalutes. These are usually count-
able on all but the smallest larvae. Additional definitive
differences between the smallest larvae of these two spe-
cies, based on plankton preserved material, remain to be
described. Blackburn (1973) indicated differences exist
in the shape of melanophores on the pectoral fin and
head, but we have not found this to be consistent and
reliable. Marliave (1975) stated that freshly preserved
larvae of P. paradoxus have more melanin in the pectoral
fins which obscures their orange color while the pectoral
fins of G. sigalutes appear bright orange. Thus there may
be real differences in the melanistic pigment patterns on
the pectoral fins of small larvae, but the limits of varia-
tion of those patterns remain to be defined. Pectoral fin
pigment differences are apparent in larger preserved
specimens with those of G. sigalutes being much darker
and having unpigmented circles near the margin.
Marliave also stated that P. paradoxus larvae develop a
notch dorsally in the caudal finfold during fin ray forma-
tion whereas G. sigalutes larvae do not. This was not ob-
vious on all specimens we have examined. Marliave
observed that P. paradoxus settles from the plankton
~13-14 mm TL while G. sigalutes remains pelagic until
>20 mm TL. Larvae of an additional form from the Gulf
of Alaska and Bering Sea, tentatively called Malaco-
cottus zonurus - Type 1 by Richardson and Bond (foot-
note 9), are also very similar in appearance to larvae of
G. sigalutes and P. paradoxus. Small larvae of all three
are nearly identical but larvae of M. zonurus - Type 1
develop 4 preopercular spines with an anteriorly di-
rected accessory spine at the base of the second spine.
This secondary spine is visible on larvae by ~7 mm upon
close examination. Pectoral fin ray counts, ~20-22, will
further separate it from G. sigalutes. Often the pelvic fins
of M. zonurus - Type 1 appear to be withdrawn into
pockets in the skin, although this condition is variable.
This form has very loose skin and remains pelagic at least
until ~24 mm. Positive identification of these larvae as
M. zonurus awaits resolution of apparent taxonomic
problems in the genus Malacocottus (Howe and Rich-
ardson footnote 3; Richardson and Bond footnote 9).
In larvae of Myoxocephalus polyacanthocephalus,
which hatch ~7 mm, melanophores cover most of the
anterior one-half to two-thirds of the body except the
ventral surface of the abdomen. The posterior one-third
of the body remains conspicuously unpigmented except
for a series of small melanophores along the ventral
midline which extends to near the tail tip. The pigment
pattern is similar to other dark cottid larvae, e.g., Radu-
linus asprellus, Rhamphocottus richardsoni,
Scorpaenichthys marmoratus, Blepsias cirrhosus, except
that a greater proportion of the tail region remains un-
pigmented laterally and the unpigmented tail region also
retains a series of ventral midline melanophores. The
head develops prominent spines including at least 4 pre-
opercular, a parietal, a nuchal, a postocular, and several
in the posttemporal-supracleithral region.
Early larvae of Orthonopias triads resemble those of
Artedius spp. with which it is closely related (Howe and
Richardson footnote 3). At hatching they have pigment
over the gut and a series of ~35 postanal ventral midline
melanophores. Soon after hatching pigment is added to
the interorbital region and on the nape. Apparently no
gut diverticula are present. Characters given in the liter-
ature are inadequate to distinguish them from Artedius
harringtoni except for possibly a higher number of ven-
tral midline melanophores.
ACKNOWLEDGMENTS
Examination of larval cottids from other areas of the
northeast Pacific was helpful in working out identifica-
tions of our Oregon cottid larvae. We thank the following
people with collections from particular areas: northern
California — David W. Rice (Lawrence Livermore Labo-
ratory); Washington and Puget Sound — Jean R. Dunn
and Kenneth D. Waldron (Northwest and Alaska Fish-
eries Center, National Marine Fisheries Service, NOAA),
Lawrence Moulton (University of Washington), James
Blackburn (Alaska Department of Fish and Game
formerly of University of Washington), Andrew Lamb
(Pacific Environment Institute, Vancouver), British Col-
umbia— W. E. Barraclough (Fisheries Research Board of
Canada, Nanaimo) and Daniel J. Faber (Canadian
Oceanographic Identification Center); Bering Sea and
Gulf of Alaska — Pat Wagner and George Mueller (Uni-
versity of Alaska). Jean R. Dunn provided reared larvae
of Hemilepidotus hemilepidotus. Kevin Howe, Oregon
State University, provided transforming and juvenile
cottids collected in Oregon tidepools and general infor-
mation on tidepool collections. James Blackburn and
Jeffrey B. Marliave, Vancouver Public Aquarium, pro-
54
vided specimens of Gilbertidia sigalutes and Psychro-
lutes paradoxus for comparative examination. Many
people helped in one way or another in working out larval
cottid identification problems particularly E. H. Ahl-
strom, NOAA, NMFS Southwest Fisheries Center;
James Blackburn; Carl E. Bond, Oregon State Uni-
versity; Jean R. Dunn; Kevin Howe; Joanne L. Laroche,
Oregon State University; Wayne L. Laroche, Oregon
State University; Kenneth Waldron; and Sharon Roe,
formerly of Oregon State University. We thank E. H.
Ahlstrom and Carl E. Bond for reviewing the manu-
script and offering helpful comments. Special thanks are
extended to Jean R. Dunn whose efforts made this paper
possible. This research was supported in part by NOAA-
NMFS Contract No. 03-78-M02-120, 1 January 1978 to
30 September 1978.
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56
NOAA TECHNICAL REPORTS
NMFS CIRCULAR AND SPECIAL SCIENTIFIC REPORT— FISHERIES
GUIDELINES FOR CONTRIBUTORS
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