430 

NOAA Technical Report NMFS Circular 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 

National Oceanic and Atmospheric Administration 

National Marine Fisheries Service 

56]1 2 
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cember 19~7. iii + 44 p., 2 figs, 1 table, 57 charts. 

408. Collection of tuna baitfish papers. (20 papers.) By Richard S 

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409. Marine flora and fauna of the northeastern United States 

Copepoda: Cyclopoids parasitic on fishes. By Ju-Shey Ho. Febru 

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415. A basis for classifying western Atlantic Sciaenidae (Teleostei: 

Perciformes). By Labbish Ning Chao. September 1978, v + 64 p., 41 

figs., 1 table. 

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1975. (20 papers.) By Julien R. Goulet, Jr. and Elizabeth D. Haynes

Editors. December 1978, iii + 350 p. 

417. Guide to the identification of genera of the fish Order 

Ophidiiformes with a tentative classification of the order. By Daniel M

Cohen and J¢rgen G. Nielsen. December 1978, vii + 72 p., 103 figs., 2 

tables. 
NOAA Technical Report NMFS Circular 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 

Juanita M. Kreps, Secretary 

National Oceanic and Atmospheric Administration 

Richard A. Frank, Administrator 

National Marine Fisheries Service 

Terry L. Leitzel!, Assistant Admlntstrator for Fisher es 
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material 

mentioned in this publication No reference shall be made to NMFS, or 

to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that N:YIFS approves, recommends 

or endorses any proprietary product or proprietary material mentivned 

herein, or which has as its purpose an intent to cause directly or indirectly 

the advertised product to be used or purchased because of this ,,'~IFS 

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 ................................................................... 17 

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 Gunther ........ ..... ..................................... 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 C linocottus 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 ofIcelinus 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 H emilepidotus spinosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 

25. Larvae of Nautichthys oculofasciatus .. .. .............................................. 41 

26. Larvae of Radulinus asprellus .................................... . . .... ..... .. . ...... 43 

27. Young of Radulinus asprellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 

111 
28. Larva of Radulinus boleoide~ ........ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lIi 

29. Young of Rhamphocottu~ richard~ont .................................................. 17 

30. Larvae of 'corpaentchth:rs marmoratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ... . 

31. Young of Scorpaenichth.vs marmoratu.\ .......•................................ ........ II 

32. Larvae of ottoid Type A .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ............ Ii~ 

Tables 

1. SpecIe of adult cottids whose range of distribution reportedly mclude marin vr bra kl h te of 

Oregon and theIr meri tic~ .................... . ....................... . :l 

2. Grouping of cot tid larvae descnb d in thi guide ba cd un preoper ular pine and plgm nt p tterns ........................................................ . 

3. l\leristics from larvae ofArtediu~ harrinRtunI ......................... . 

4. lea urements (mm) o/Iarvae oL4.rtediu harrmgtonl .. . . . ......... . 

Meristics from larvae ofArtedlu., Type 2 ...... ...... . . . ............. . 

6. leasurements (mm) of larvae 01 Artediu '1 ype 2 ....................... . 

7. Men tic from young of Clinocuttu . acutlcep ...... ... . ............... . 

8. 

9. 

10. 

11. 

12 . 

13. 

14. 

15. 

16. 

17. 

18 

19 

20 

21. 

22. 

23. 

24. 

25. 

26. 

27. 

28. 

29. 

30. 

31. 

32. 

33. 

34. 

35. 

36. 

37. 

38. 

39. 

40. 

41. 

42. 

43. 

44. 

45. 

Measurements (mm) of larvae 0/ C linocottu acutlcep .. ............... . 

Menstics of larvae of Cottidae Type 1 •...•.•.•........•.•. . ••.•..• 

Mea urement (mm) of larvae 01 uttidae Type 1 ••.••••• • ••••••• • •.•• 

Meri~tics from larvae of l uttidae '[ ype 2 ...... ...... . . . ........ . 

Mea 'urements (mm) ot larv e of 'ottidae T 'pe 2 .,. . . . . . .. ....... . ... . 

• 1eri tICS from larvae ofCottidae Type 1 ............... . ............. . 

l\lea urements (mm) oflarvae of ottidae '1 ype 3 ................ . 

:\leristics from larvae 0/ TnRlops sp. .. ................... .... . ... . 

l\leasurement (mm) oflan ae 0/ Tnglop p. ............. ...... . .. 

Meri tics from larvae of ChltoTlotu., pugetensi . . . . . . . . . . . ... ..... . 

Measurements (mm) ollarvae of Chltonotu pugett!nsi ................... . .. 

Menstics from larvae of Cottu., a per ..................................... . 

Mea urement (mm) of larvae of Cottus a.~per . . .. ...................... ..... . ...... . 

Menstics from larvae of Enophn ., bi~on ........ ...................... . ... . 

Measurements (mm) 01 larvae of EnophT) bison ................................ ..... . 

l\leristic from larvae of Irelznu spp ................................................... . 

Mea urements (mm) oflarvae of Icelmu' pp..... . ..................................... . 

Meri tics from Ian ae of Leptocottus armatu ......................... . ............... . 

Measurement (mm) of larvae 0/ Leptocottus armatus .............................. . . . 

Meristic from young of Pancelinus hopliticu ..................... ... ............... . .. 

Measurements (mm) of young of Pancelinu . hopliticu,!; ................................... . 

Meristics from young of Hemilepidotu . hemilepidotus .................................... . 

Measurements (mm) of young Hemilepidotus hemilepidotu ............................... . 

Meristics from young of Hemilepidotu~ spinosus . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . ....... . 

Measurements (mm) of young of Hemilepidotu ' Sp!TlO~U .................................. . 

Meristics from larvae of Nautichthy oculofasciatus ...................................... . 

Measurements (mm) of larvae of Nautichthys oculofasciatu ............................... . 

Meristlcs from young of Radulinus asprellus ............................................ . 

Measurements (mm) of young of Radulinus asprellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . .. 

Meristics from larvae of Radulinus boleoides . . . . . . .. . .................................. . 

Measurements (mm) of larvae of Raduhnus boleotdes ..................................... . 

Meristics from larvae of Rhamphocottus richard Onl ...................................... . 

Measurements (mm) of larvae of Rhamphocottus richardoni ............................... . 

Meristics from young of Scorpaenichthys marmoratus .................................... . 

Measurements (mm) of young of Scorpaenichthys marmoratu ............................. . 

Meristics from larvae of Cottoid Type A ................................................ . 

Measurements (mm) of larvae of Cottoid Type A ........................................ . 

Additional northeast Pacific cottid species for which larvae are known ........................ . 

IV 

1 

5 

5 

1 

15 

15 

23 

24 

24 

~4 

.,-

~O 

~7 

3 

31 

32 

32 

32 

33 

34 

36 

40 

41 

42 

42 

42 

45 

45 

45 

46 

46 

50 

50 

51 

52 

53 
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 summarized. 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. 

Lan'ae of Artedius harringtoni, Chitonotus pugetensis, Clinocottus acuticeps, Icelinus spp., 

Paricelinus hopiiticus, 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 armat;"s, Nautichthys oculofasciatus, 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 cottid larvae include morphology, pigmentation, head spination, and meristics. 

Literature summaries are also provided for northeast Pacific cottid larvae in 8 additional genera: 

Biepsias, Das)'cottus, Gilbertidia, ?Malacocottus, Myoxocephalus, Oligocottus, Orthonopias, 

Ps),chroiutes. 

INTRODUCTION 

The cottids are one of the most speciose and diverse 

oups of fishes inhabiting coastal waters of the northst Pacific Ocean with 90 species in 40 genera reported 

occur from Baja California to the Aleutian Islands 

Howe and Richardson l). The range of 36 of these 

ecies in 20 genera reportedly includes waters off 

regon (Table 1). Adult cottids range in size from the 5 

Artedius meanyi, Clinocottus acuticeps, and 

ligocottus rimensis to the 76 cm Myoxocephalus polycanthoceihalus and Scorpaenichthys marmoratus 

art 1973). Larger forms in the genera Gymnoanthus, Hemilepidotus, Hemitripterus, Melletes, and 

yoxocephalus are fished commercially by the 

.S.S.R. and Japan (Gorbonova 1964; Fedorov 1973). 

ff Oregon several species are taken frequently by 

ports fishermen, e.g., Enophrys bison, Hemilepidotus 

emilepidotus, H. spinosus, Leptocottus armatus, Scor-

~aenichthys marmoratus (Beardsley and Bond 1970). 

'School of Oceanography, Oregon State University, Corvallis, Oreg.; 

present address: Gulf Coast Research Laboratory, East Beach Drive, 

cean Springs, MS 39564. 

'School of Oceanography, Oregon State University, Corvallis, OR 

97330. 

Howe, K., and S. L. Richardson. 1978. Taxonomic review and 

I eristic variation in marine sculpins (Osteichthys: Cottidae) of the 

rortheast Pacific Ocean. Final Report, NOAA NMFS Contract No. 03-

178-M02-120, 1 January 1978 to 30 September 1978, 142 p. Northwest 

~nd Alaska Fisheries Center, National Marine Fisheries Service, NOAA, 

~725 Montlake Blvd. East, Seattle, WA 98102. 

1 

Most, if not all cottid larvae hatch from demersal eggs 

-(about}1-4 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 larvae 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 1 V2 yr 

survey off Oregon they occurred in 80% of the samples 

but comprised only 8'b 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 difficult because of the large number of species in the family, 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 Col!. Prog. Pub!. No. 

ORESU-T-77-003, 73 p. 
tu acuticeps, Rhamphocottus richardsoni). Small «6 

mm) cottid larvae have a very characteri tically protruding snout which, with development, can vary considerably in its degree of pointedne (e.g., compare 

Scorpaenichthys marmoratus and Radulinus asprellus). 

Several form have unusual gut diverticula or protrusions from the abdominal wall (Artedius Type 2, 

Clinocottu acuticeps) which serve a triking 

characters. 

Pigment patterns can range from darkly pigmented 

form uch a Scorpaenichthys marmuratus and 

Radulinus asprellus where melanophore cover the entire 

body except the tail tip to relatively unpigmented form 

such a Triglops sp. with melanophore only on the head 

and gut. In many specie (Artedius harnngtoni. 

Chitnnotu pugetensLs, Paricelmu hopliticus) the 

number. extent, pacing, and hape of the potanal 

ventral midline melanophores erve a diagno tic aid . 

even though these may decrea e in number with development. Ranges and frequency of ount of the e melanoph ores within developmental period are useful. The 

presence or ab ence of nape pigment and/or head pigment, particularly in mall «5 mm) larvae. may be 

helpful in orne ca es (Artediuli pp .. Clinocottu' pp , 

Enophrys pp.). The pattern and extent of pigment over 

the gut is u eful in otherwi e light I} pigmented form . It 

may range from relatively heavy pigment over the entire 

gut (Paricelinus hoplitLCUS. Tnglop ' p .) to relativel} 

light concentrations over the dorsolateralurface only 

(Cottidae Type 1, Cotbdae Type 2).ometlme it appears in distinctive patterns (the bar m Leptocottus armatus, the distinctive round melanophore in Enophry 

bison) although these patterns can be obscured in poorly 

or improperly pre erved material. 

Most cot tid larvae develop four spine on the preopercleo However, some forms develop only one prominent 

spine (Rhamphocottus richardsoni) while other develop 

a large number, up to ~25, of smaller spines (Artedlus 

spp.) on the preopercular margin. A ingle parietal and 

nuchal spine may develop (lcelinu pp.) or a clu ter of 

smaller spines (Cottidae Type 1) or no spine at all (Leptocottus armatus). The same situation may prevail in the 

posttemporal-supracleithral region. One or more postocular spines may develop (Hemilepidotus spp., Triglop 

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 

4 

counts for a particular . p 'cies ar' generally 4 Of Ie. ". 'I til 

ddT rence between dOT. al (U) and anal (A) olt IfJ) 

counts may abo b of help which i u ually D ./ A, <:.g. 

by :3 to 5 rays in Artpdius pp., Cllnocullu pp., anc 

()1t~(Jc{Jltu.\ spp. and >9 in authlchth) (JcuilJ/a cwtu 

In some sp cie the lJ und A count are nearly equa 

(Lep/oc(Jttw) and in ome lJ (Punceiinu hfJplLll 

cus). 

Oth r characters that may be u elul include ile 

hatchmg, sIze at tran iormali(}n, and cirri pattern. rIb 

size at hatching, e.g., -3 mm in ArtF'dlU harrmRtOnL v 

~l~ mm In Hemltrlpteru am rlcanu (War el and Mer 

I1man 1944' Oklvama and ando 1976) and the Ize a

transforma;ion, ~.g., -9 mm m Enophf) bl on and -If 

mm in Chitunotu pugeten I, cnn erve a dl 

lmgui. hmg character. He d mi pallern-, which rna, 

be eVident in older larvae, m 1y be Important in linkin 

larvae .... 11 h known juv nde . How ver, they appear t( 

develop after lh pelagl ph e in R number of p cie, 

e.g., heltrlU pp., and may b f limited value. 

ORDER Of PRI<.. Ll 'lATIC .. 

I (I ta dltate u e ot thl gUld, th larvae de cnbe 

herein have been grouped a ordmg to ertam prim 

churacte", n able 2). H(J .... \er, a formal dichotomou 

ke} 1. not pr)\ ided man} pe I remain to be de 

cnbed and u hake} ould b mi leadin . Rather 

group. are t()rmed on th b 1 of the m tout tandin 

Ian al character. I he two major groupin are ba ed 0 

preopercul r pine pattern: ~r JUp . numerou (>4 

group B, 4 or Ie . \\ llhm group ,the larvae may b 

lurther di. tlngul. hed b) u mg a combin lion ot charac 

teL includincr number of pr opercular pine, number 0 

ventral midlIne melanophore. number ot parieta 

pine . and pre ence or abo ence ot gut diverticulae 

Within group B the larvae may be placed in four ub 

groups on the ba i of pu tanal pigmentation. Withi 

the e lour group. larvae can be further eparated u in 

characters a for group A and relative body hape 

pigmentation. and number of myomere, 

Artediu harringtoni ( tark 

(Figures I. 2; 1 able a. 1) 

LIterature A 4.2 mm "'L larva re-embling A. harrin 

tom wa de cribed by Blackburn (1973) a Cottid 6. Lac 

vae of thi pecie were reterred to a Artedlu p. 1 b 

Richardson ( ee footnote 4), and Rlchard on and Pearc 

(1977). 

Distinguishing Features. Larvae of A. harringtoni are 

probably 3-4 mm long at hatching. The large t 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. 

A. Preopercular spines numerous (>4) 

Artedius harringtoni 

Artedius Type 2 

Clinocottus acuticeps 

Cottidae Type 1 

Cottidae Type 2 

Cottidae Type 3 

B. Preopercular spines 4 or fewer 

1. Postanal pigment absent 

Triglops sp. 

2. Postanal pigment on ventral midline only 

Chitonotus pugetensis 

Cottus asper 

Enophrys bison 

Icelinus spp. 

Leptocottus armatus 

Paricelinus hopliticus 

3. Postanal pigment primarily on dorsal 

and ventral midline on small larvae 

Hemilepidotus hemilepidotus 

Hemilepidotus spinosus 

4. Postanal pigment heavy over body 

Nautichthys oculofasciatus 

Radulinus asprellus 

Rodulinus boleoides 

Rhamphocottus richardsoni 

Scorpaenichthys marmoratus 

Cottoid Type A 

' Maximum number at peak of development. 

Preopercular' 

spines 

11-21 

13-23 

8-13 

19 

10-11 

16-19 

4 

4 

4 

4 

4 

4 

4 

4 

4 

4 

4 

4 

o 

' From Howe & Richardson 1978 (see text footnote 3). 

Ventral' 

midline 

melanoph ores 

27-30 

14-24 

7-9 

7 

15-17 

4-5 

o 

41-45 

14-17 

11-14 

10-11 

9-11 

31·32 

14 

24-34 

N3 

N 

N 

N 

N 

N 

Parietal' 

spines 

0 

Multiple 

0 

Multiple 

1 

Multiple 

1 

o 

1 

o 

1 

B 

o 

Vertebrae' 

(myomeres) 

32-34 

(32-34) 

32-33 

(32-35) 

(32-34) 

(33) 

(47) 

35-36 

34-39 

29-31 

33-39 

35-39 

42 

35-31 

35-37 

40-41 

38-39 

39-40 

26-28 

35-37 

35 

3N = not distinguishable; B = bump. 

Additional 

distinguishing characters 

dorsal gut diverticula 

hindgut diverticula 

hindgut bulges 

dark gut 

deep bodied 

finfold pigment 

pigment bars on gut 

dark gut 

dorsal pigment interrupted 

dorsal pigment continuous 

large pectoral fins 

slender bodied 

deep bodied 

globose 

Table 3.-Meristics 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 ra:is 

Anal Pectoral fin Preoper- midline Dorsal Ventral Vertebrae Branchi· 

Body Dorsal fin fin _ fin ra:is 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 30 N N N N N N N 

--------------------------- ------------------------------- --------------------------------------------------

7.3 -17-** 13 14 14 16 26 N N N N N N 7 

9.3 IX 16 13 14 14 I,3 21 22 N N N N N N 7 

11.2* IX 17 14 14 14 I,3 21 N 5 6 6 3 11 23 7 

13.6 IX 17 13 15 15 I,3 18 23 N N N N N N 7 

13.6* IX 17 13 14 14 I,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 notochord flexion.) 

Body 

length 

3.0 

4.7 

6.9 

7.3 

9.3 

13.6 

Head 

length 

0.66 

0.82 

1.8 

2.0 

2.0 

4.3 

Snout 

length 

0.16 

0.20 

0.39 

0.47 

0.35 

1.1 

Eye 

diameter 

0.32 

0.39 

0.58 

0.78 

0.94 

1.1 

Snout to 

anus length 

1.3 

1.6 

3.3 

3.4 

4.5 

6.6 

Body depth 

at pectoral 

fin base 

0.72 

1.1 

1.6 

2.4 

3.0 

3.8 

'Multiple spines develop along preopercular margin instead of usual 4. 

5 

Pectoral 

fin length 

0.12 

0.31 

0.47 

0.90 

2.0 

3.6 

2d preopercular 

spine length' 

N' 

N 

N 

N 

N 

N 

Parietal 

spine length 

' N = Not measured. 
Arfed/us harr/ngfon/ 

3.0mm NL 

4 .7mm NL 

t ....... ~~! ... ~\. 

----------~=--

6.9mm NL 

Figure I.-Larvae of Artedius harringtoni. 

6 
Arfed/us horr/ngton/ 

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 characteristically extend as dashes of pigment out onto the finfold 

particularly in larvae >7 mm. Some pigment is added to 

7 

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 larvae. 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 pecimens, possibly due to fusion or overgro\vth by ti ue. Artedius harringtoni never develops 

pronounced multiple spines in the parietal and posttemporal region as does Artedius Type 2 although minute serrations are sometimes visible and a bump 

develop in the parietal region in larger specimens. 

Larvae of A. harringtoni are stubby in shape and oft en 

have a humped appearance in the nape region. The 

hindgut trails below the body. Snout to anus length is 

generally <50co 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 33C'0 SL. The length of the pectoral fin increases from 4 to 26CO SL during larval development. 

Artedws harringtoni is the only species of Artedius with 

evpn branchiostegal rays (Howe and Richardson footnote 3) even branchiostegals were countable in alllarval speClmens examined (25) down to ~7 mm. The innermost pair of branchiostegals is extremely fine and small 

In relatior 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 

DiscussIOn. Identification of this series as A. harring toni was based primarily on meristic characters, especially the presence of seven branchiostegal rays. 

Relative abundance of adults in the area of larval capture 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 

(Howe 6). 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. harringtoni. Within these three species, only A. harrington! 

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 spin 

while A. lateralis has 1 or 2 cirri at the base of each spine 

The largest specimens in our series have a single nasa 

cirrus at the base of each spine and the largest specime 

has small bumps in the preorbital region which may b 

developing cirri. 

Artedius Type 2 

(Figures 3, 4; Tahles 5,6) 

Literature. Blackburn (1973) briefly described and 

illustrated (8.5 mm SL) larvae resembling our Artedius 

6K. Howe, Ph.D. candidate, Department of Fisheries and Wildlife

Oregon State University, Corvallis, OR 97331, pers. commun. Septembe 

1978. • 

Table 5.-Meristics from larvae of Artedius Type 2. (Specimen between dashed line is undergoing notochord flexion.) 

peclmen stained with Alizarin Red S; •• = Dorsal fin spines and soft rays cannot be distinguished ; B = Bases only; N = Not 

examined.] 

Pelvic Ventral Caudal fi n a~s 

Anal r ·ctoral fin Preoper- midline Dorsal Ventral Vertebrae Branchi- Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostega! length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays 

.l.U N 24 N N N N N N N ~ - ;-.! -5 14 N N N N N N N --------------------------------------- -- -----------------.------------------.-------

IiO :\' 13 -15 N N N N N N N --------- ------- --- --------------- --------------------- -----------------------------

14H-" 12B N 21 15 N N N N N N 6 CJ.C) \ III 12 15 N 1,3 23 18 N N N N N N 6 

I S' 1:\ Hl 12 15 15 1,3 24 N 6 4 12 22 6 .' , 1:\ 17 1;~ 15 K 1,3 13 13 N N N N N 6 

"Iea~urements (mm) of larvae of Artedius Type 2. ( pecimen between dashed line is undergoing notochord 

n exion .) 

Hod) 

Itngth 

o 

Head 

length 

0.66 

12 

Snout 

length 

U.18 

0.12 

Eye Snout to 

diameter anus length 

0.30 1.2 

U.44 1.9 

Body depth 2d preoperat pectoral Pectoral cular Parietal 

fin base fin length spine lengt h spine length' 

0.58 0.22 N3 N 

1.2 0.20 N N 

------------------------------------------------------ l II 1 i 

20 

o 

11 41 

•• J!) 0.56 

0. 1 U.66 

o ~" 0.94 

D.t>-! 113 

2.8 1.5 

-------------- --------------

3.5 2.1 

4.6 2. 

5.6 3.0 

~llllt.r e plm dt'HI"p altlfll( preopercular margin Instead of usual 4. 

lult,p pille- ut'\elop In regIOn of pane tal spine 

I t me ur I 

0.31 N N 

--------------- ------------------

0.86 N 

1.7 N 

3.4 N N 
Artedius Type 2 

3.0mm NL 

3.0mm NL 

------~ 

; .. / .-/~ 

4.7·mm 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 

1 

J 

... . - . 7 2mm SL 

1 

K - ..... r-r99mm SL 

~~ A -- . rf.@' 

\£: 1 9) .' .... " ' • I ~~... !. . 'II .::: ... 

. .!:, ~ 

, \', .:... A, t';' -~ 

.. 

II 8mm SL 

Figure I.- Lan ae of Ar/ediu,; T\ pe 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 Cottid No.4. White (1977) illustrated (3.9 mm NL) and 

briefly described similar larvae as Cottid ill 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 resemble 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 

10 

(1975) illu trated and briefly de cribed young tage (4

8. 11. 14 mm TL (total length)) of.4.. LateralllS He al 

di cu sed egg of.4. lateraliiS and egg and Ian ae of .4. 

fenelStralis Larvae of thi form were called ArtedlU p. 

by Richardson (footnote 4) and Richard on and Pearcy 

(1977) . 

Distinguishing Features. Larvae of ArtedlUs Type 

probably hatch at around 3-4 mm. The largest specimen

we have captured pelagic ally are -14 mm, taken in 

neuston net 2 km off ewport, Oreg. They are beginnin

to develop juvenile pigmentation. 
In larvae, melanophores are concentrated over the 

orsolateral surface of the gut. Nape pigment is usually 

esent in larvae except at the smallest sizes but 

ecomes embedded and obscured in later stages > 8 mm. 

row of usually <20 (often 15-20 but <25 in larvae <5 

m) melanophores occurs along the ventral body 

idline posterior to the anus. The number of ventral 

tidline melanophores helps distinguish Artedius Type 2 

om A. harringtoni which usually has >20. Approxiately the posterior half of these melanophores extend 

5 characteristic dashes of pigment onto the ventral finld especially in larvae >7 mm. In the largest 

pecimens, ~12-14 mm, pigment is added to the head 

egion, at the base of the pectoral fin, to the anterior porn of the spinous dorsal fin, as a bar extending from the 

rsal surface of the abdominal cavity near the dorsal 

rigin of the pectoral fin to the middle of the spinous doral fin, and as groups of melanophores along the lateral 

Ilidline from the gut region to the caudal fin. 

As in A. harringtoni a series of small spines develops 

long the margin of the preopercle, beginning to appear 

4.5-5.0 mm. Tbey increase in number to ~20-25 in 

ostflexion larvae, ~7-1l mm, then decrease in the largt specimens. Those at the dorsal edge and at the posrior angle of the preopercle become large relative to the 

st of the spines. Clusters of spines develop in the 

larietal region beginning with 1 or 2 spines in 6-7 mm 

arvae, 3 spines in 7.5-8.5 mm larvae, and more on larger 

pecimens. By ~9 -9 .5 mm clusters of spines occur in 

~oth the parietal and posttemporal regions, and these renain visible on the largest specimens examined. Such 

pine clusters are never obvious on A. harringtoni. 

Larvae of Artedius Type 2 are similar in shape to A. 

wrringtoni being stubby and often with a hump in the 

~ape region. The hindgut appears to trail below the 

Dody. Snout to anus length is <50% SL. A prominent 

reature 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. haringtoni, are most pronounced in small larvae, becoming 

less conspicuous as the pectoral fin develops. They are 

present on the largest pelagic specimens we examined. 

his larval character is probably lost during transformation 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 

Cievelopment. Branchiostegal rays, which could be 

ounteq on specimens >7 mm, consistently numbered 6 

or all larvae in this series compared to 7 for A. haringtoni. 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 collections seem more indicative of A. fenestralis than A. 

_____ 11 

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. harringtoni, 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 occipital 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 insertion 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, presence 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 (Ahlstrom 7) 

beyond the southern range limit of Diablo Cove, Calif., 

reported for A. fenestralis (Miller and Lea 1972). 

However, A. lateralis does occur there. Final confirmation 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 collections. 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. 
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. 

C//nocottus acuticeps 

3.7 mm NL 

3.7mm NL 

". 

3.9mm NL 

12 
. Figure 6. 

Clmoco . ffus acuficeps 

-Young 0 

7.6mm SL ' 

f Clinocottus acuticeps. 

1~ 

16.5mm SL 
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 probably 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. Pigment is also present on the head between the eyes in all 

but the smallest specimens which may be faded. Melanophores 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 melanophores are on the ventral body margin anterior to that 

point. Melanophores in this ventral midline series are 

rather inconspicuous compared to other species. The posterior melanophores in this ventral row do not extend as 

dashes onto the finfold as in Artedius spp. but may actually occur on the ventral finfold rather than on the 

body margin in larger larvae. A single melanophore persists 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 transformation, pigment is added to the head, pectoral fin base, and 

onto the anterior end of the spinous dor al 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 decreases during transformation with only 1 dorsal spine 

vi ible in a 19.4 mm stained juvenile. 0 additional head 

spine are obvious in larvae. Parietal and nuchal spines 

are never apparent although the flabby skin could posSibly obscure their pre ence on unstained larvae. 

Larvae of C. acuticeps have an unusual gut, the posterior portion of which trails well below the body. Near the 

anus two diverticula or protrusion are evident throughout 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 enca ed in a bubble of 

100 e outer skin, particularly in the head region. Body 

depth ranges from -22 to 33cc L during larval developTable 7.-Meristics from young of ClinocottuB 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 = Ba e only, D = 

Damaged; N = Not examined I 

Pelvic Ventral Caudal fin rays 

Anal Pectoral fin Preoper- midline Dorsal Ventral Vertebrae BranchtBody Dorsal fin fin fin ra~s spine cular melano- econd- Prin- Prin- econd- Abdom- Cau- ostegal 

length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays 

3.7 N 7 ~ ~ ~ '\ :\ ~ :\ 

3.9 N 9 ~ ~ :\ ~ :\ :\ :\ 

------------------ -------- ------------------------ --------------------------------------------- -----._------

6.9 -22B-** 11B 14 11 8 N N N N N ~ :\ ---------- ---------- -- -------------------------- ---------- -------------------------------------------------

7.6 VII 15 11 14 N Buds 11 7 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 14 N 1,3 13 7 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 N D D D D 10 23 6 

Table B.-Measurements (mm) of larvae of Clinocottus acuticeps. (Specimen between dashed line is undergoing notochord flexion.) 

Body depth 2d preoperBody Head Snout Eye Snout to at pectoral Pectoral cular Parietal 

length length length diameter anus length fin base fin length spine length ' spine length 

3.7 0.90 0.14 0.38 2.3 0.90 0.26 N' 

3.9 0.88 0.16 0.38 2.4 0.86 0.40 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 

4.2 

0.47 

0.58 

0.86 

0.70 

0.94 

1.2 

5.4 

7.0 

9.0 

2.5 

2.6 

4.2 

'Multiple spines develop along preopercular margin instead of usual 4. 

14 

1.2 

1.6 

4.1 

N 

N 

N 

= Not measured. 
ent. The adult complement of fin rays (or elements) 

an be counted in larvae by ~7-8 mm. Late stage larvae 

ave relatively long pectoral fins, ~30% SL. On larger 

rvae, ~13-14 mm, the inner pelvic fin ray is noticeably 

ttached to the belly by a membrane. This diagnostic 

haracter helped link the larval series to identified 

iscussion. Larvae of C. acuticeps are similar to 

tlescribed larvae of C. analis and C. recalvus in size at 

atching, ~3-4 mm, and presence of nape and head pigent and pigment over the gut on small larvae . (The 

ack of head pigment on the smallest larvae of C. 

cuticeps may have been caused by fading.) All three 

species have a series of ventral midline melanophores but 

hose on C. analis and C. recalvus appear to be higher in 

number than C. acuticeps. The posterior melanophores 

n this series do not appear to extend out onto the finfold 

. n C. acuticeps as reported for the other two species. 

ased 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, mainly 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. lC by Richardson (footnote 4) and Richardson and 

Pearcy (1977). 

Distinguishing Features. The smallest larvae in our collections 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 unpigmented. Melanophores 

occur at the nape, in a rather light cattering over the 

dorsolateral gut surface and in a row of < 10 along the 

ventral midline posterior to the anu . The e ventral 

midline melanophores are well spaced beginning -4-5 

myomeres behind the anus and extending to the po teriormost myomere. They decrease in number anteriorly 

with development. 

The preopercle develops a high number, -19, of mall 

spines on its margin. A cluster of tiny spine develop in 

the parietal region, a few of which are visible on 4 mm 

specimens. Another cluster of minute spine develop 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 ~23-26cc SL. The 

snout is relatively round in appearance. 

Discussion. Specific or generic identification of Cottidae Type 1 is not possible without additional material. 

These larvae are related to the Artedius-Clinocottus-Oligottus 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 appear to most closely resemble Artedius spp. The cluster 

of spines in the parietal and posttemporal- upracleithral region in these larvae have been observed only 

in Artedius Type 2 and Cottidae Type 3. They do not occur in C. acuticeps, C. recalvus (Morris 1951), or O. 

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 I. (Specimen between dashed lines is undergoing notochord flexion.) 

\ entral 

Dorsal Dorsal Anal Pectoral Pelvic fm Preoper- m,dltne 

Body fin fir fin fin rays spine cular melanolength spines rays rays Left Right and rays pine phore, 

4.2 N' 

6.7 19 

9.0 IX 18 12 15 ]\i Buds 19 3 

'N = Not examined. 

Table 10.- Measurements (mm) of larvae of Cottidae Type I. (Specimen between dahed lines is undergoing nolo hord 

fl exion. ) 

Body depth 

Body Head Snout Eye Snout to at pectoral 

length length length diameter anus length fin base 

4.2 0.84 0.20 0.42 1.8 0.98 

6.7 1.4 0.20 0.52 2.6 1.6 

9.0 2.1 0.25 0.94 4.2 2.5 

I Multiple pines de\·elop along preopercular margin instead of usual 4. 

' Multiple pines develop in region of parietal spine. 

' N = Not measured. 

15 

~d preoper 

Pectoral euler 

fm length 

0.20 

0.55 

12 
Cotlldo lyp 

..... 

• • .. ' . " " , 

" 42mm L 

67 m L 

9 Omm SL 

II un'i. I In" of ( ollld I I pr I 

larger pecimens to determine pooitl\eiy if the pattern i 

the same as in our two Artediu . type!', 'I hi preopeH ul r 

spme pattern differs trom that in C acutjcep and (' 

recalvu~ (Moms 19.'51) in which the dur:-ulmo t spine 

becomes the large t. The number of preopercular ~pine 

in Type 1 is relatively high, ~ 18 as in A harringtonj and 

Artedius Type 2 (>20), compared with the lower 

numbers for C acuticepli (l0-15), C. recalvu (4·8), and 

O. maculosus (7-9). The gut shape and length resemble 

our ArtedLUs types. 

Based on the meristic compilation by Howe and 

Richardson (footnote 3) (Table I), count on our large t 

Cottidae Type 1 specimen, particularly the 18 dorsal soft 

16 

n t 

not f) maculo u 

(1. -2, 1 73) I lentlfi 

tern twm our Type 1, and one would e peet pattern 

!'imdar to () nlllculo.,u, on O. :myderl. Problem. In iden· 

tifying A it?T1t?,truil\ were discu ,,.ed under ..\rtedju, Type 

:2 and cannot be re 'olved at thi: time. 
Cottidae Type 2 

(Figure 8; Tables 11, 12) 

sp. 20 by Richardson (footnote 4) and Richardson and 

Pearcy (1977). 

'Literature. Larvae of this type have not been 

reviously described. These larvae were called Cottidae 

Distinguishing Features. The smallest larvae in the 

series are ~4 mm and recently hatched. The largest 

specimen is 7.4 mm in which the bases of dorsal and anal 

Cattidae Type 2 

4.0mm NL 

6.4mm NL 

···. A . .. ~_____ ____________~--------

··;;·······.·~.:.\ ····:;··············> ----- ------- ·· \) ~.:.~ ..... . 

. -- \ - , 

::=,0i_SJ -'. Md. 

-- ,- /------' 

~-- 7.4mm NL 

Figure B.- Larvae of Cottidae Type 2. 

17 
fin rays are just forming. Size at transformation is unknown. 

Preflexion larvae have pigment at the nape, a relatively light scattering of melanophores over the dorsolateral surface of the gut and a series of <20 but.> 10 

ventral midline melanophores po terior 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 la t myomere. Some melanophores in this series radiate out onto the finfold. A few 

melanophores occur on the finfold near the tail tip. With 

development some melanophore' are added to the head 

and the number of ventral midline melanophore decreases. Melanophores were not apparent on the caudal 

fin base of the largest pecimen, a would be expected 

based on the pigmentation near the tail tip of mailer larvae, but the specimen wa badly faded. 

A series of approximately 10 mall pines develop on 

the preopercular margin. The upper spine appear omewhat thicker than the lower pine on the large t 

specimen. A tiny parietal pine i vi ible on the 7.4 mm 

specimen but no additional head region pine are apparent. 

The shape of the gut is rather di tinctl\'e with the 

hindgut trailing noticeabl) below the body, similar to 

Clinocottus acuticep . Snout to anu length i around 

50l" SL, longer than most pecies. When viewed ventrally, the wall of the hindgut bulge out on each ide of 

the anus remini cent of the hindgut diverticula of C. 

acuticeps but not as pronounced. Body depth at the pectoral fill base is ~21-27(o L. The nOut i quite rounded. 

The larvae have a flabby appearance, particularly 

around the head reglOn, imilar to C. acuticeps. 

Discussion. The multiple preopercular pine pattern of 

Cottidae Type 2 indicates thi type is related to the Artedius-Clinocottus-Oligocottus group. Ba ed on a combination of larval characters it seems to re emble 

Table II.-Meristics from larvae of Cottidae T)pe 2. ( pecimen~ 

below dashed line are undergoing notochord flexion.) 

Body 

length 

4.0 

6.4 

7.4 

Dorsal 

fin 

spines 

Dorsal Anal Pectoral 

fm fin fin rays 

rays rays Left Right 

Pelvic fm 

spine 

and rays 

Preopereular 

spines 

10 

11 

Ventral 

midline 

melanophores 

15 

17 

6 

Ciinocottu\ most closely. It hu. a relillivel leJw number 

«15) of multiple spine on the prerJpf!rcle 11 drJ!: ( . 

acutiCf'PS (10-1.'») und C. r('calLius (G-9) (MeJfrl 19'=:;1). 

The upper prcopercular pine appear tf) be enlargin~ 

relative to the re t, a~ Jll (' acult(;('ps and C. rpcaivu 

Only a tiny panetal pin IS vi ible on the large t 

specimen. Two (Iny pinE'. appear in the parietal and 

nuchal po. i(lons in (' rr'ca/tu but only for a brie! period 

and none were 011 erved in ('. acutlcrp. pos ibly r)b 

s('urcd by thC' nabby kin. ,\lelanrJphore appear on th 

napl' and head of all ('. TI'caiuu and all but the malle,t 

C. aw/lceps nel ( ottidae '1 ype 2. The lack of pigment 

on the head of our malle t peclmen could be are. ult a 

lading. ( iin()clJtI u rrcaluu Inrvac were de cnbed from 

rcarl·d and tre,hly pre en'cd muterial which often teneL 

to hu\' increa. ed numb r ul and more pronounced 

melan()phore . The hindgut trail. well below the body a .. 

in C. awl/rep,. Bulge appear in the hindgut wall on 

cach SIde of thc anu"imilar to the more pronounced 

dl\'erticula ell ( . acutlcep uch protuberance were not 

not d for C. rr calt;u , perhap on ob ervational over-

. ight. :ome melanophore in the ventral midline erie 

radiate ont() the finlold a In ( • recaluu . Thi quality 

wa not n()ted in C' flcutlcep but po, ibly could be 

related to quality of pre ervatioll. ~lelanophore occur on 

the lin fold ncar the tail lip, often 8.1 treak of pigment in 

all three form!'>. '1 he,.,e melanophore. later appear on the 

caudal fin in '. a uttC p and ( . recaluu '. They were not 

\'i Ible em our larget ottidae Type.!. pecimen po~ ibly 

becau e It wa. faded. 

ottidae Type :3 

(Figure 9; Table~ l:l. III 

Literature Lan'ae of- thi type have not been 

pre\'iou Iy de cnbed. They were Ii ted by Richard n 

(footnote 4) and Ri hard on and Pearcy (1977) as 

Oltgocottu.~ -po l. 

Di.~tingw 'htng Feature. The malle t larvae ''Ie have 

taken are -5 mm and relatively undeveloped. Our 

large t pecimen i 12,5 mm. ~10 t fin appear fully 

formed except for econdary caudal ray. but final transformation mu t occur at a omewhat larger ize. 

The e larvae have inten e pigment over the head and 

nape even in the malle t pecimen . 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 

Body depth 2d preoper- Head Snout Eye Snout to at pectoral Pectoral cular Parietal length length length diameter anus length fin base fin length spine length' spine length 

4.0 0.8 0.16 0.42 2.2 0.84 0.24 N' ----------------------------- ---------------------------- ---------------------- ------------------------ 6.4 

7.4 

1.5 

1.8 

0.27 

0.55 

0.55 

0.66 

3.0 

3,9 

1.6 

2.0 

'Multiple spines develop along preopercular mar"';n instead of usual 4 2 N = Not measured. "'. 

18 

0,27 

0.66 

N 

N 0.10 
Cottidae Type 3 

6.3mm NL 

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 intense and densely concentrated, appearing as large round 

melanophores. The only other pigment consists of ~4-5 

ventral midline melanophores near the posterior most 

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 

19 

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 C linocottus 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 accomplished with certainty. Based on current knowledge 
of larval characters, particularly the multiple preopercular spine pattern, it belongs with the Artedius-Clinocottus-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. recaluus, and O. maculosus which all have 

<15 (Morris 1951; Stein 1973). The upper spine in Cottidae 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:l. (Sp('c-imcn. 

above dashed line are undergoing notochord 0 ·XlOn.) 

Ventral 

Dorsal Dorsal Anal Pectoral PelVIC fin Prcoper- midline 

Body (in fin fin fin ray" spine cular molanl) 

length spines rays rays Left Right and rays pines ph'Jrcs 

6.3 7 5 

7.5 19 4 

12.5 IX 15 1:2 1-1 14 1,3 16 

Table 14.-Measurements (mm) of larvae of Collidae Type 3. (Specimen., above da,hed line arc undergoing notochord flexion .) 

Body Head Snout Eye 

length length length diameter 

6.3 1.4 0.31 0.62 

7.5 1.9 0.58 0.74 

Body depth 

Snout to at pectoral 

anus length fin ha,e 

3.0 1.4 

4.2 ~.:! 

Pectoral 

{In length 

0.27 

0.56 

:ld preoper 

cular 

pine len!1h 

~ 

Parietal 

pine length 

---------------------------------------------_ ... 

12.5 3.4 0.80 1.1 7.:3 :U 

'Multiple spines develop along preopercular margin instead of usual !. 

'Muillple spines develop in region of panetal spIne. 

IN = Not measured. 

Type 1 and unlike C. acuticeps, C. recaluus, and O. 

maculosus. However, spines in the parietal cluster appear 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 posttemporalsupracleithral 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 Cottidae 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 Oligocottus 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 Clinocottus species. 

Triglops sp. 

(Figure 10; Tables 1.5, 16) 

LIterature, Larvae identified a Triglops p., which 

resemble oUI larvae, were dIu trated (8.3, 12 mm L) by 

Blackburn (1973) but no descriptive information was 

given. Larvae of other pecies of '!'riglops which have 

been de cribed include 10-18 mm T. ptngeli (Ehrenbaum 1905-1909; Koefoed 1907 [cited by Khan 1972]; 

Ra3 1949) from the Atlantic Ocean and Barent ea and 

8.4, 11.6, 18.9, 23.4 mm TL T. murrayi (Khan 1972) from 

the Atlantic Ocean. 

Distinguishing Features. Our mallest preflexion larva 

is 6.9 mm and relatively undeveloped. The largest 

Table 15.- Meristics from larvae of Trig/ops sp. (Specimens aboH 

dashed line are preOexion, that below is postflexion.) 

Ventral 

Dorsal Dorsal Anal Pectoral Pelvic fIn Freoper- midline 

Body fIn fin fIn fin rays spine euler melanolength 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 i. 

postflexion. ) 

Body depth 2d preoperBody Head Snout Eye Snout to at pectoral Pectoral cular Parietal 

length length length diameter anus length fin base fIn length spine length s pine length 

6.9 1.9 0.20 0.90 2.6 1.5 0.39 

7.4 1.8 0.20 0.86 3.1 1.4 0.39 

15.4 4.4 0.80 1.4 6.8 3.2 2.9 0.62 BRI 

'BR = Broken. 

20 
Triq/ops sp. 

\ 

/~~ ) --- r "--" 

6.9mm NL 

15.4mm SL 

Figure lO.-Larvae of Triglops sp. 

specimen we have is 15.4 mm. It has a large complement 

f 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 pectoral fin base. The rest of the body remains unpigmented. 

Head spines on the largest specimen include a double 

row (anterior and posterior series) of 4 pre opercular 

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 pectoral 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 distinguish 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 northern waters off Washington where two species of Triglops 

reportedly occur, T. macellus and T. pingeli (Hart 1973). 

21 

But coastal currents generally flow northward during 

that time of year, minimizing the likelihood that the larvae 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 complements 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 discussion 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 
o the cond dorsal fin 

mm and 8 medlol ter ) row by 11 mm 

I ed b) Kh nIl:.?) ) If lan"ae of T. 

m the P clfic de\elop th me pl;m :1t path r m the tlantlc nd Barent I:a they 

11\ dl tm~1 h ble from our Tn~ ·r ,'p.lard nd trunk plg'ment and hape of T. pin~ell 

r mb ur Tn lop -p mdl ting' a eneric imilarity. 

Chitonotus pugetensis ( teindachner) 

(Figures 11 , 12 ; Tables 17. IS) 

Litert>:'urt: Lan ae of thl 

pre\"lOU Iy de cnbed. 

pecie - haye not be 

Fi~ur" II.-Lan ae of Ch iton otus pUKc ('n,~is. 

Chi/ono/us puqe/ensis 

r : • 

. . .,~.~ 

... ... ~ ..... -

3.0 mm NL 

6 .3mm NL 

•••• 0'· 

....... 

.... _ • 0- .' . 

8.5m m SL 
Chitonotus pugetens/s 

Figure 12.-Y oun g of Chitonotus pugetensis. 

. (Speci- . I vae of Chitonotus pugetensts. fl xion.) T bl 

17 -MeristICs from ar . those below are post e a e . r e preflexlOn, mens above dashed me ar Ventral 

Pectoral Pelvic midline Dorsal Dorsal Anal fin Preoper- melano- spine cular 

fin fin fin rays spines phares Body fin 

length 

3.0 

6.3 

spines rays rays Left Right and rays 45 

8.5 -15-' ~: 16 16 

11.5 IX ~~ 16 18 18 1,3 

4 

4 

15.4 X 15 17 17 I,3 

16.6 X 15 t be distinguished, 

4 

4 

'Dorsal fin spines an d s aft rays canna 

41 

37 

24 

26 

24 

23 

llest larvae we have . , F atures. The sma . h t h d The Distingwshmg e d robably recently ~ c e. _ 

taken are ~3 mm ~n p 6 6 mm is beginnmg to tra?s 

largest pelagic spe~lme~, Ie . as evidenced by increasmg form to the juvemle s ag 

t

· . I moderately pigmenta IOn. f the gut IS on y 

The dorsolateral surfac~ 0 t distinctively lines the 

' ented. Additional plg~en I 'ty Pigment occurs plgm h bdomma caVI . , ventral margin of tea d' s added to the head regIOn 

on the head by 8.5 mm an 1 A series of melanophores 

through later development: ostanally. These ventral 

lines the ventral body margm ~umber > 20 and u~ually 'dline melanophores alway

6

s This is the hlgh~st 

~~O in preflexion la~va e < f::~phores for preflexlOn num b er of ventral mldlme me 
Ta ble lB.-Measurements (mm) of la rvae of Chilonotus pugetcnsis. (. peciml'ns a bove dab hed line ure prcl1 ('x ivn. lh oM' 

below a rc postl1exion.) 

--- - Body depth 2d pr~(Jp~r 

Body Head Snout Eye Snout to at pectoral Pectoral {"lJlar Parietal 

length length length diameter anus length fin base 

3.0 0.66 0.23 0.70 0.54 1.1 

6.3 1.7 0.43 0.62 2.6 1.5 

8.5 2.2 0.72 0.80 4.0 2.0 

11.5 3.5 0.88 1.1 5.3 2.8 

15.4 4.6 1.1 1.6 7.4 3.8 

16.6 6.2 1.8 1.7 7.6 3.9 

IBR = Broken. 

larvae of any species described in this guide. The posterior 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 anterior 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 durmg 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 Icelinus spp. where the body outline narrows distinctively 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 between 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 . 

lin len!(th spi~..!:ngl~ ~p~n~ I~~gth 

0.20 

0.31 

0.64 0.12 1).04 

1 6 023 HH 

2.6 0.31 (J 12 

4.0 0.40 

Cottus asper Richardson 

(Figure I:J; Tab e~ 19,20) 

LIterature. Larvae of Cattus asper were briefly de -

cribed and illustrated (5.5, 9.0, 10.8 mm TL) by tein 

(1972). We dId not review literature on larvae of tnctly 

freshwaterpecie of ('attus. 

DLstlnguishLng Features Larvae hatch -5 mm and 

begin to develop juvenile pigmentation -10 mm a evidenced by an Illcrease in the number of melanophore in 

the head region. 

Cottus asper larvae are relatively lightly pigmented. A 

few melanophore - cover the dor al urface of the gut and 

everal elongate melanophores line the ventral margin ot 

the abdominal ca\'ity and throat. About 15-20 evenly 

spaced melanophores occur along the ventral mIdline of 

the tail beginning near the anu . The e decrease in 

number during development while a few melanophore 

are added laterally over the gut and one appears at the 

ventral edge of the preopercle. As tran formation begins 

more pigment is added to the head, ba e of pectoral fill, 

over the gut, and along the base of the caudal fill. 

Table 19.-l"l eristies from lanse of Cotlus asper. (Specimen a bo,-e 

dashed line is preflexion. those belo" are postflexion.) 

Body 

length 

5.2 

8.2 

9.9 

Dorsal Dorsal Anal Pectoral 

tin fin fin fin rays 

spines rays rays Left Right 

:\'-

-19-' 16B' :\ 

IX 18 16 19 18 

IN = Not examined. 

Peine fm 

spine 

and rays 

Preopereular 

spines 

4 

4 

'Dorsal fin spines and soft rays cannot be distinguished. 

'B = Bases only. 

Ventral 

midline 

melanophores 

E 

14 

8 

Table 20.-Measurements (mm) of lar vae of Cottus asper. (Specimen a bove dashed line is prefl exion , those below a re 

postflexion. ) 

Body Head 

length length 

5.2 0.9 

------------------

8.2 

9.9 

1.9 

2.6 

Body depth 2d preoperSnout Eye Snout to at pectoral Pectoral eular Parietal 

length diameter anus length fm base fin length spine length spine length 

0.12 0.36 2.1 0.78 0.51 

---------------- - -------- ----------------------------------------------------- - ----

0.35 

0.66 

0.55 

0.74 

24 

3.8 

4.2 

1.4 

1.8 

0.62 

2.5 

-0.03 

0.33 
Col Ius asper 

5.2mm NL 

~.---.-~ 

,<,. \ 

I 

/ ... . . J .. :: "' ~ 'f 4. )t.).......¥ ,. A . .,. "" j 

....- --- .- -. ---. '------~-..... -.-. _____ .J----r-,./ 

S.2mm SL 

Figure 13.-Larvae of Cottus asper. 

Four pronounced spines develop on the pre opercular 

argin, evident in 8 mm larvae. No additional head 

pines form with development. 

The shape of C. asper larvae is rather distinctive, being 

omewhat 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 plankton collections. RingstadB 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 hisutch}: an experimental and ecological study in a British Columbia 

coa tal stream. Environ. Can., Fish. Mar. Serv., Tech. Rep. 457, 88 p. 

25 

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 generally 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 deTable 21.-Meristics from larvae of Enophrys bison. (Specimen between dashed lines is undergoing notochord flexion.) 

Ventral 

Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline 

Body fin fm fin fin rays spme cular 

length spines rays rays Left Right and rays 

4.8 :-.; ' 14 

7.0 4 11 

9.1 LX 11 9 16 1,3 10 

'N = Not examined. 
Enophys bison 

4.8mm NL 

......... .,. "~'1 ").. 

.). ~ 

OJ. .c· 

Figure 14.-Young of Enophrys bison. 

26 

7.0mm NL 

--;'?/3~¢c) < - . . ..... -. £~, 

""" ... c:-._.;_ ~ 

9.lmm SL 
Table 22.-Measurements (mm) of larvae of Enophrys bison. (Specimen between dashed line is undergoing notochord 

flexion.) 

Body Head Snout Eye Snout to 

length length length diameter anus length 

4.8 1.1 0.2 0.55 2.3 

7.0 2.0 0.5 0.66 3.8 

9.1 3.4 0.6 0.90 5.0 

scribed a 7.5 mm larva of E. bison and Marliave (1975) a 

10 mm TL larva. Misitano (1978) described a developmental 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 laboratory but Marliave (1975) captured a 10 mm TL transforming specimen in the plankton and we took transforming 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 material but are faded on our smallest specimens. Melanophores are present on the nape, appearing very dark in 

freshly preserved specimens. The number of post anal 

ventral midline melanophores is usually ~15. A distinctive space of several myomeres in length exists 

between the anus and the anteriormost ventral melanophore. The ventral midline melanophore series is relatively 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 

spinO\~s 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 extending 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 

Body depth 2d preoper· 

at pectoral Pectoral cular Parietal 

27 

fin base fin length spine length spine length 

1.1 0.37 

1.9 0.55 0.35 0.27 

3.4 2.08 0.43 0.27 

nasal spine appears on the snout. Two spiny tips (midopercular 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 larvae <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 transformation. Enophrys bison larvae have a characteristically low number of myomeres (vertebrae = 29-31) 

(Howe and Richardson footnote 3) which helps to distinguish 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 likely 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 melanophore, 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. lilljeborgi (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 Richardson (footnote 4) and Richardson and Pearcy (1977). 

Distinguishing Features. The smallest larvae in our collections are ~3-4 mm and probably recently hatched. 

The largest specimens we have collected pelagically are 
feel/nus spp. 

3.3mm NL 

8.6mm NL 

IO.9mm SL 

.- _ .. ~" ... , 

...~ '-: .. 

<' •• 

J-:. 

t 

, .... 

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 dorsally 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 distincti ve 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 

28 

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 specimens, > 12 mm, have 2 distinct melanophores on the dorsal 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 specimens, with and without dorsal midline melanophores, 

have an additional distinct pigment spot on the dorsalmost 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. 
leel/nus spp. 

13.8mm SL 

~ 

~: 

-'-~ 

15 2mm SL 

... - ... -... ~.,. 

16.5mm 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 qevelop and fuse together. Several spines appear 

in the posttemporal-supracleithral region eventually being 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 large t specimens. 

The gut shape is rather di tinctive in larvae of Icelinus pp. Snout to anu length is relatively short in 

mall larvae, <40""0 "L, but increases to -500(. L on 

larger specimens. Body depth at the pectoral fin base is 

~20 ( SL in small larvae and ~25-30cc "L on older larvae. Larvae of Icelinus spp. have a very characteri tic 

29 

body shape. Body depth appears to be con tricted ju t 

behind the anus, bulge out lightly in the mid tail re~ion. 

and then narrow di tinctively again near the tail tip or 

caudal peduncle. They al 0 develop a rather polllted 

nout which i apparent on larvae> mm. Pectoral fin 

length is -25-30' "'L on the large. t pecimen .. 1 he 

adult complement of fin ray or element i. countable on 

pecimen > 12 mm including the diagnu tic pelVIC fin 

count of 1,2. 
/eel/nus spp. 

12.5mm 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 fm rays 

Anal Pectoral fin Preoper- midline Dorsal Ventral Vertebrae BranchiBody 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 10 N N N N N N N 

5.1 11 N N N N N N N 

----------------------------------------------------------------------------------------------- ------------. 

8.6 3 10 N N N N N 

----------- --------------------------------------------------- --------------------------------------- --------

10.9 -25-** 12 15 15 4 

t 12.5 X 16 12 16 16 1,2 4 

13.8 X 16 12 16 16 1,2 4 

t1·U X 16 12 16 16 1,2 4 

14..1' XI 15 12 15 15 1,2 4 

15.2 X 16 12 15 15 1.2 4 

ti5.5* X 16 12 15 15 1,2 4 

16.5 X 16 12 16 15 1,2 4 

t 16.6 X 16 13 15 15 1.2 4 

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, 1. 

burchami, 1. filamentosus, 1. oculatus, and 1. tenuis 

overlap. An anal fin ray count of 12 agrees only with 1. 

burchami from Oregon waters (Table 1), a species which 

IS reportedly rare (Howe and Richardson footnote 3). 

Four other northeast Pacific species of Icelinus (1. 

boreahl:i, 1. cavifrons, 1. fimbriatus, and 1. quadri10 

8 

10 

11 

11 

N 

10 

12 

30 

N N N N N N N 

N N N N N N N 

N N N N N N N 

N N N N N N 

5 6 6 4 1l 23 6 

N N N N N N N 

6 6 6 4 11 23 6 

N N N N N N N 

N N N N N N N 

seriatus) can have an anal ray count of 12 but none 

reportedly occur off Oregon (Howe and Richardson footnote 3). Characters used to distinguish adults such as cirri patterns and elongated dorsal fin spines are not 

developed on our largest pelagic specimens. These factors together with inability to find consistent larval characters to split our Icelinus spp. larvae into subgroups potentially equivalent to species forces us to keep our 

indentification at the generic level. 
Table 24.-Measurements (mm) of larvae of Icelinus spp. (Specimen between dashed line is undergoing notochord 

flexion.) 

Body Head Snout Eye Snout to 

length length length diameter anus length 

3.3 0.76 0.10 0.32 1.2 

5.1 0.86 0.12 0.39 1.7 

8.6 1.8 0.51 0.55 3.4 

10.9 2.5 0.80 0.88 4.6 

112.5 4.0 1.0 1.2 6.2 

13.8 4.2 1.0 1.3 6.4 

114.1 5.2 1.4 1.4 7.0 

15.2 5.2 1.2 1.5 7.7 

16.5 7.2 2.7 1.4 9.0 

t16.6 6.1 1.8 1.4 8.3 

tForm with 2 melanophores on dorsal midline. 

Leptocottus armatus Girard 

(Figure 18; Tables 25, 26) 

Literature. Eggs and recently hatched larvae (~4 mm 

Body depth 2d preoper· 

at pectoral Pectoral cular Parietal 

fin base fin length spine length spine length 

0.68 0.26 

1.0 0.31 

1.7 0.39 

2.2 1.04 0.22 0.12 

3.0 2.2 0.32 0.26 

3.6 2.4 0.55 0.31 

4.2 3.0 0.48 0.24 

4.0 4.1 0.56 0.24 

4.8 3.8 0.56 0.24 

4.2 3.2 0.32 0.32 

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 l8.-Larvae of Leptocottus armatus. 

LepfocoftU5 armatu5 

----~ ------~ . '. . . ... ,. " "'---."" 

~~ ~ \ ---------~ ~ 5.lmm NL 

8.lmm NL 

Il.lmm SL 

31 
Tabl :!5.-;\t~li from lanae of Leptocottus armatus. (pecimen 

betwe<!n da hed line i undergoing notochord flexion.) 

Bod) 

length 

51 

- 1 

III 

Don;al DoC'<8.1 Anal Pectoral 

fin fin fin fin ra ''; 

ray~ 

15 

'Ill Ii 19 

ot examined. 

Ventral 

Pelvic fin Preoper. midline 

cular melano· 

pines phores 

11 

9 

4 7 

Discussion. Leptocottus is a monotypic genus in the 

northwest Pacific. The larvae with their characteristic 

bars of pigment over t he 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 t he snout serves as a useful 

diagnostic character. 

ble 2r..-.\tea.urement., (mm) of laT\'ae of Leptocottus armatus. (pecimen between dashed lines is undergoing 

notochord flexion.) 

Body depth 2d preoper· 

Hody H~ad , 

nout Eye noutto at pectoral Pectoral cular Parietal 

l~ngth length length diameter anus length fill base fin length pine length spine length 

5.1 0.92 0.1 0.25 2.0 0.84 0.23 

1.9 031 0.32 3.2 1.6 0.78 0.10 

111 2.. o.62. ___ .::::o.::::82:..-__ ___.:4~.9~ __ ___.:2::..3=__ ___ _=.:2 . .::.2 _____ 0.::.: .::.21:...... _____ _ 

(lH77) illu. trated (4.9 mm NL) and briefly described 

mall larvae. Blackburn's Cottid 5 (5.8 mm SL) was 

probably thl pecies as was Eldridge's (1970) Cottid No. 

1 (51 mm). 

Dl tmgw htng Features Hatching takes place at 3.9-

. mm '1 L. Our large. t pelagic specimen is 11 mm and 

m,} be nearing transformation as indicated by incrca cd pigmentation over the head. Marliave's (1975) 13 

mm TL .peeimen \\as transformed and had assumed a 

b nthlC h bit. 

Larv e have distInctive pigment over the dorsolateral 

urf e t the gut \\ hieh always appears as ~6-8 bars exc pt m damaged pecimen·. 1 '0 other known cottid lar· 

\ h \e uch gut pigment. Larvae al 0 have distinctive 

Illt rn I l)lgm nt on the. nout at the anterior part of the 

f r bram and xtending po;;teriorly from the eye to the 

p toml fin Th lutter becomes obscured by muscle but 

th former remam vi ible throughout the larval period. 

I' I m I nophorc occur along the ventral margin of 

ut cit. \ I' '" of u uall ' < 15 rather prominent 

n n phor 0 cur along th \ entral midline of the tail 

nmn bout 3 m,}omer~ p )~terior to the anus. \Vith 

I pill nt. m lfln phore are added to the head and a 

t the ba of the caudal tin. 

L 

32 

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 evidenced by increased head pigmentation and body spination (modified scales). A 25.6 mm specimen captured 

pelagically appears to be a fully transformed juvenile. 

Larvae have a distinctively pigmented gut with melanophores covering the entire abdominal cavity. Pigment 

is cattered 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 decreases to -15-20 in postflexion stages. At the posterior 

end of this ventral row a separate group of distinctive 

melanophore occurs near the tail tip extending onto the 

finfold. The e melanophores line the base of the caudal 

tin after it forms. Little pigment is added during the larTable 27.- Meristics from young of Paricelinus hopiilicus. (Speci' 

ml'n, abow dashed line are preflexion. those below are postflexion.) 

DONal Dursal Anal l'~doral 

Hod~ fin fin lin fln ra 

length ~plne:-. rny~ ray., Len Right 

5.0 ' 

f" ). 

- .---- .... _- ..... __ ..... _- -- XlI lH 23 15 

• '11 HI ~3 I'> 

. 'II I~ 23 If> N 

- • ot t' lirDlne<! 

Pelvic fin 

.pine 

and ray~ 

1,5 

1,5 

1,5 

Preoper. 

ular 

8plne~ 

~ 

I 

4 

Ventral 

mldhne 

melano· 

phores 

32 

31 

17 

17 

19 
Table 28.-Measurements (mm) of young of Paricelinus hopliticus. (Specimens above dashed line are preflexion, those 

below are postflexion.) 

Body 

length 

5.6 

6.2 

13.8 

18.6 

25.6 

Head 

length 

1.2 

1.7 

4.3 

5.0 

8.6 

'BR = Broken. 

Snout 

length 

0.23 

0.31 

1.0 

0.96 

2.1 

Eye 

diameter 

0.51 

0.70 

1.7 

1.4 

2.7 

Snout to 

anus length 

2.0 

2.2 

7.0 

8.8 

11.5 

al period except over the head and at the base of the 

pectoral fin. The pelagically captured juvenile has much 

'ncreased pigmentation in blotches over the entire body 

nd 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 

piny scales that develop dorsally. Similarly, several 

Body depth 

at pectoral 

fin base 

0.86 

1.0 

4.2 

4.6 

5.3 

Pectoral 

fin length 

0.16 

0.23 

BR' 

4.2 

7.4 

2d preopercular 

spine length 

BR 

0.80 

0.96 

Parietal 

spine length 

BR 

-0.24 

0.48 

spines appear to develop in the posttemporal-supracleithral 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, inPar/eel/nus hoplit/eus 

5.6mm NL 

6.2mm NL 

Figure 19.-Larvae of Paricelinus hopliticus. 

33 
Par/eel/nus hoplit/eus 

18.6mm SL 

25.6mm SL 

Figure 20.-Young of Paricelinus hopliticus. 

creasing to ~25-30% SL during the larval period then decreasing in the juvenile stage. The general body shape of 

the larvae resembles that of Icelinus spp. The snout becomes 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. 

Hemilepitodus hemilepidotus (Tilesius) 

(Figures 21,22; Tables 29, 30) 

Literature. Gorbunova (1964) illustrated (7.25, 10.5 

mm) and briefly described larvae of H. hemilepidotus. In 

Table 29.-Meristics from young of Hemilepidotus hemilepidotus. 

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). 

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 specimens as large as 23 mm have been captured pelagically 

(Specimen between dashed lines is undergoing notochord flexion.) 

[* = Specimen stained with Alizarin Red S; N = Not examined.] 

Ventral Caudal fin ra):'s Scales 

Anal Pectoral Pelvic fin Preoper- midline Dorsal Ventral Vertebrae Branchi- Dorsal above 

Body Dorsal fin fin fm rals spine cular melano· Second- Prin- Prin- Second· Abdom- Cau- ostegal scale lateral line 

length Spines Rays rays Left Right and rays spines phores ary cipal cipal ary inal dal rays rows Left Right 

5.8 N 14 N N N N N N N N 

5.9 N 14 N N N N N N N N 

9.1 N 2 9 N N N N N N N N _________________________________________ _____________________________ 'l.. ____________________________________ __ ________ • _____ 

10.7 IX 19 15 17 N Buds 4 11 N N N N N N N N 

11.4' 17 12 13 13 Buds 4 N 6 6 12 19 6 

11.5 XI 18 15 17 N Buds 4 7 N N N N N N N N 

19.0 XI 20 16 16 16 1,4 4 2 N N N N N N N 4 19 N 

23.0' XI 19 17 16 16 1,4 4 N 7 6 6 6 12 24 6 5 27 23 

34 
t the surface. Thus there may be a pelagic juvenile 

hase similar to that of rockfishes, Sebastes spp. The 

'mallest transformed specimen collected in Oregon tideools was 21 mm SL. In our collection we have no specimens 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 

he posterior 3d or 4th myomere. An unpigmented area 

ccurs on the dorsal midline between about myomeres 4-

n which fills in with pigment by ~6-7 mm. The dorsoateral surface of the gut is moderately pigmented. A 

series of melanophores extend along the ventral midline 

beginning at about the 9th or 10th po tanal myome 

extending to all but the last 3 or 4 myomere . i\lelanophores in this series usually number < 15. With development, melanophores are added over the head, anterodorsal body surface, and the gut. orne internal 

melanophores appear to extend dorsally from the ventral midline melanophores in larvae as small as 6 mm. 

Additional internal melanophores appear ventrolaterally between the notochord and ventral midline. By 9 

mm, internal melanophores can be seen in a row above 

Hem//epldofu5 hem//ep/dofus 

5.8mm NL 

". ~ , 

Figure 21.-Larvae of Hemilepidotus hemilepidorus. 

35 

5.9mm SL 

., . 

9 .l mm NL 
Ta ble 30.-:\Ieasurements (mm) of young of Hemilepidotus h emilepidotus. (Specimen between dashed lines is undergoing notochord flexion .) 

Body depth 2d preoperBody Head nout Eye Snout to at pectoral Pectoral cular Parietal 

length length length diameter anus length fin base fm length s pine length spine length 

5. 0.90 0.06 0.52 2.0 1.0 0.32 

5.9 1.0 0.27 0.58 2.1 0.86 0.34 

9 1 2.0 0.27 0.9 3.4 1.7 0.62 0.22 0.10 

10.7 2.7 0.47 1.2 4.6 2.6 1.1 0.39 0.27 

11.5 3.7 0.96 1.3 6.4 3.3 1.7 0.51 BRI 

19.0 6.7 1.28 2.2 11.0 6.0 4.0 0.9 0.31 

BR = Broken. 

Hem//ep/dotus heml!epldotus 

IO.7mm SL 

11.5mm SL 

19.0mm SL 

Figure 22. Young of Hcmiiepidiltus hemilepidotus. 

36 
nd below the notochord along much of the body. These 

nternal melanophores remain visible until the lateral 

usculature obscures them. External melanophores are' 

,ot added laterally until larvae are.> 11 mm, and then as 

I relatively light scattering compared to H. spinosus. 

he ventral midline melanophores gradually become inernal but remain visible through the skin and serve as a 

. iagnostic character to distinguish them from H. spinous until dorsal scale rows develop (see Discussion). By 

9 mm the entire body except the caudal peduncle and 

entral body margin are covered with pigment which is 

I arker dorsally and lighter below. 

Four prominent spines develop on the preopercle, some 

I fwhich are visible by 9 mm. A parietal spine can be seen 

it ~9 mm and later a nuchal spine, the two fusing to-

~ether late in the larval period. A postocular spine 

[evelops over the eye ~9 mm and remains obvious at 

east until 11 mm, probably disappearing during trans-

'()fmation. This spine develops only as a minute spinule 

'or a brief period, ~8-9.3 mm in H. spinosus, barely visiDie without staining. With development three spines appear in the posttemporal-supracleithral region, being 

reduced to bumps in larger specimens. A cleithral spine 

ts 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 

y ~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 

[9 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 

FY ~14 mm. Gill membranes are noticeably free from the 

Isthmus in the 19 mm specimen. 

is cuss ion, The relative position of the first postanal 

entral midline melanophore and the posterior extent of 

the ventral series serve as excellent diagnostic characters to separate larvae of H. hemilepidotus from H. spinasus until the dorsal scale rows are formed. In H. hem iepidotus 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-

~ra l row begins immediately behind the anus and conEists of usually > 15 melanophores in all but the largest 

iopecimens. In larger specimens, the number of dorsal 

scale rows separate the two species, 4 or 5 in H. hemilepidotus and 6-8 in H. spinosus. These can be counted in 

specimens as small as 14 mm. 

The placement and number of ventral midline melanophores 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 

37 

do not appear to be adequate to distinguish larvae among 

these four species. One character used by Gorbunova to 

distinguish late larvae of H. 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. hemilepidotus, and Peden (1964) has shown them in young of H. jordani and H. zapus. These become overgrown on larger 

fish. Gorbunova stated that larvae of H. gilberti develop 

5 pre opercular 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 together with a more detailed examination of externallateral 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 described by Gorbunova (1964) are easily distinguished by 

the extension of the dorsal and ventral midline melanoph ores 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 under H. hemilepidotus.) 

Distinguishing Features. The smallest larvae in our collections are ~5 mm and appear to be recently hatched. 

Specimens as large as 27 mm have been captured pelagically 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 posterior most myomere. 

Some pigment occurs on the dorsolateral surface of the 

gut. A row of > 15 melanophores extends along the ventral 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 hypural elements. With development external melanoph ores 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 noticeable in a line above the notochord along most of the 
Heml/ep/dolus sp/nosus 

5.0mm NL 

--- ---- ' .. ~-~ 

__ ~. ----.1,- • ~ 

---------- 8.9mm NL 

Figure 23.-Larvae of Hemilepidotu., spino.us. 

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 melanophores become embedded but remain visible through the 

skin at least to -14 min, which serve to distinguish these 

larvae from H. hemilepidotus. 

38 

Four preopercular pine' develop, beginning -7 mm. 

A parietal pine can be een -7 mm and later a nuchal 

spine develop posterior to it fu mg with it in the largest 

specimens. A minute po tocular 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-
Hemi/ep/dotus spmosus 

II.Omm SL 

11.8mm 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. 

')Q 

spinosus are much more abundant in our plankton samples 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 common. Collections in Oregon tidepools have yielded primarily young juveniles of H. hemilepidotus and only 

rarely H. spinosus. Our data indicate that H. hemilepidotus generally spawns closer to shore and the larvae are 

not as neustonic as H. spinosus. Tidepools may be important nursery areas for juvenile H. hemilepidotus whereas 
T b 31 ~ll'ri li~ from ~oun of Hemilf?pidotus spino us. (_pl?'Cimen between dashed lines is undergoing notochord flexion.) 

• • I zann Red ... . .. ; D",-.al rID spines and oft ray" cannot be dlstmgui ' hed; B = Bases only. :\ = :\ot examined. I 

Yemral 

Anal P ora 

Caudal fin ra'.::\·_~~_ 'cales 

Peh,c fm Preoper- mIdline --Dorsal \'ertebrae Branch!- Dorsal above 

fin fm ra} , ' pme cular melano- :erond- Prin- 'econd- Abdom- Cau- ostegal scale laterallin€ 

ra) Le Ri hI aDd ra) ,pines phore' ary i:!: al ip~aI __ an=.::.. _· _ __ iD-,-a_I _ _ d_a_l _ _ ra-=-y_s __ ro_\\ __ Le_f_t _Ri _',,-gh-! 

14B 

B·· liB 

A·· 14B 

1~ 

6 

1 " '5 

14 

1 ~ 15 

1'; 15 

1- 15 

15 

I~ 15 

.I 15 

1- 15 

Ib 1 

I" 15 

I' II' 15 

15 

IJ 

I 

I' 

I' 

6 

I~ 

1~ 

'5 

1.1 

Buds 

Buds 

Hud, 

1.4 

1,4 

Hu(b 

1.4 

1 4 

1.4 

Aud, 

1. 1 

I 4 

1 4 

I 4 

14 

I j 

I j 

1 

14 

~ 

~ 

3 

3 

4 

3 

4 

4 

4 

4 

4 

4 

4 

4 

4 

I 

j 

4 

24 

31 

:\ 

26 

, 

21 

22 

25 

21 

22 

18 

:\ 

:\ 

~ 

19 

24 

21 

22 

13 

pmo Ll utilIz e difterent areas, perhaps farh r , 

\ (lIltichthys oculofasciatu . (Girard) 

( I I u r 2.i; 'I hit"~ 3:\, :1 1) 

d 0 \ r th latud I pedun I , 

6 

6 

6 

'\ 

N 

'\ 

:-... 

6 

:\ 

6 

" 6 

3 

N 

N 

N 

N 

6 

N 

3 

5 

7 

N 

N 

N 

N 

7 

7 

:\' 

N 

12 

N 

N 

12 

12 

N 

N 

N 

N 

12 

N 

12 

12 

12 

N 

N 

N 

N 

12 

N 

N 

12 

:-; 

12 

N 

N 

1 

t\ 

N 

21 

N 

24 

N 

N 

N 

24 

24 

24 

24 

N 

N 

24 

N 

24 

1\1 

24 

:\ 

t\ 

N 

N 

N 

6 

N 

N 

6 

6 

N 

N 

N 

6 

6 

6 

N 

N 

N 

6 

N 

N 

6 

N 

6 

N 

N 

t\ 

t\ 

N 

N 

N 

N 

N 

N 

4 32 32 

6 -30 

N 

N 

6 29 N 

6 31 33 

6 24 N 

6 N 29 

6 26 26 

6 21 

6 34 30 

with melanophore extending onto the dorsal and ana 

fin fold and fin in the middle of the tail region. Melanophore also occur over the head, at the base of the pectoral fin, on the pectoral fin ray as a band near the fin 

margin. and over the dor olateral surface of the gut. Pigment i · added to the spinou dor al fin as the anterior 

rays become elongate. 

Fuur spines develop on the preopercle appearing a 

bumps. Enlarged bump' also develop in the parietal 

region. anterior to it along the parietal ridge, in the 

postocular regiun. nasal regIOn, and several in the po tt mporal-supradeithral area .• one of these appear as 

pronounced sharp spines. 

The length 01 the pelvic lins is stnking, 42 and 59', L 

in our two specimens :nout to anu length i relatively 

long, 56 nd 51 (( SL while body depth at th pectoral fin 

bll e i relatively. lender, ~1 and 23 L Th number 01 

m 'om r ( > 4U) i relatively high for a cottid. The larvae 

hsv nn a umdh °e ppearance. 

Ih cu 10ri OnlY!Jne peel ot Nautichthy.\ cur:. olf 
Table 32.-Measurements (mm) of young of Hemilepidotus spinosus. (Specimens between dashed lines are undergoing 

notochord flexion. ) 

Body depth 2d preoperBody 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 BR' 

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. 

Noutichthys ocu/ofosciotus 

11.7mm NL 

_ J,>' • • 16.6mm SL 

Figure 25.-Larvae of Nautichthys oculofasciatus. 

AJ 
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. 

pribilouius and N. robustus are unknown. 

Table 33.-Meristics from larvae of Nautichthys oculofasciatus. 

(Specimen above dashed line is undergoing notochord flexion.) 

Ventral 

Dorsal Dorsal Anal Pectoral Pelvic fm Preoper- midline 

Body fin fin fin fin rays spine cular melanolength spines rays rays Left Right and rays spines phores 

11.7 VII 28 19 13 12 N' 

16.6 IX 28 19 14 

'Not present as a distinct row. 

'N = Not examined. 

N 4 N 

Radulinus asprellus Gilbert 

(Figures 26, 27; Tables 35, 36) 

Literature. Larvae of this species have not been previously 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 characteristics 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 

11.7 

16.6 

Head 

length 

3.4 

4.5 

Snout 

length 

0.80 

1.1 

Eye 

diameter 

1.1 

1.3 

Snout to 

anus length 

6.6 

8.4 

Body depth 

at pectoral 

fin base 

2.5 

3.8 

Pectoral 

fin length 

4.9 

9.8 

2d preopercular 

spine length 

0.20 

Parietal 

spine length 

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.] 

Anal Pectoral 

Body Dorsal fin fin fm rays 

length Spines Rays rays Left Right 

4.7 

5.2 

5.9' 

6.1 

6.7 

7.2 

7.8' 

7.9 

8.5' 

9.1 

9.6 

10.1 

10.9 

-22B-*' 

VII 21 

-21-** 

X 22 

18B 

22 

22 18 

22 19 

N 

N 

Pelvic 

fin 

spine 

& rays 

Buds 

Ventral Caudal fm rays 

Preoper- midline'" Dorsal Ventral Vertebrae Branchicular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal 

spines phores ary cipal cipal ary inal dal rays 

N 

3 

N 

N 

3 

4 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N ----------- -------------- -- - _.- - -- - -- - -- - - - -- -- --- - - - ----- --- - -- - - - - -- -- ---- - - -- - ---. -- --- - - -- - - -- -- ---- ---. - - -- -- 11.4 

11.4* 

11.5 

11.7* 

12.0 

12.0' 

12.6 

12.6 

13.0 

13.0' 

13.9' 

14.1 

14.1 

14.4 

14.4 

14.7" 

15.2' 

-23-*' 

X 21 

IX 22 

X 22 

IX 22 

X 21 

IX 20 

X 23 

IX 22 

VIII D 

X 22 

IX 23 

X 22 

IX 22 

X 23 

IX 

IX 

22 

23 

22 

23 

24 

22 

23 

24 

22 

24 

24 

23 

23 

23 

23 

24 

24 

23 

24 

18 

18 

16 

17 

18 

19 

19 

18 

18 

18 

18 

19 

18 

19 

19 

18 

18 

18 

N 

17 

19 

18 

19 

19 

18 

18 

18 

19 

18 

19 

19 

18 

Buds 

Buds 

Buds 

1,3 

1,3 

1,3 

Buds 

1,3 

1,3 

1,3 

1,3 

1,3 

1,3 

1,3 

1,3 

4 

N 

4 

N 

4 

N 

4 

4 

4 

N 

N 

4 

4 

4 

4 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

3 

N 

N 

N 

N 

3 

3 

N 

5 

N 

5 

N 

6 

N 

N 

N 

6 

6 

N 

N 

N 

N 

6 

6 

N 

5 

N 

5 

N 

6 

N 

N 

N 

6 

6 

N 

N 

N 

N 

6 

6 

N 

N 

N 

N 

N 

N 

2 

N 

N 

N 

N 

2 

2 

N 

N 

12 

N 

12 

N 

N 

N 

12 

12 

N 

N 

N 

N 

12 

12 

N 

N 

20 

N 

23 

N 

N 

N 

27 

27 

N 

N 

N 

N 

27 

27 

N 

N 

6 

N 

6 

N 

N 

N 

6 

6 

N 

N· 

N 

N 

6 

6 
Radul/nus asprellus 

7.9mm 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 dorsolaterally 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 

midline§>. 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. asprelIus from Scorpaenichthys marmoratus which they resemble at small sizes. Pigmentation generally increases with 

development with only the tail tip or caudal peduncle remaining unpigmented. The lateral midline pigment is 

noticeable on all but the largest specimens. 

Four small spines develop along the preopercular 

43 

margin, visible by ~1O-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 relatively 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 complement 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 
Rodu//nus ospre//us 

IO.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 pigment differences as given in that section, at least for the 

size range described. Larvae of the third species, R. 

uinculus, 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. 

44 

Radulinus boleoides Gilbert 

(Figure 28; Tables 37,38) 

Literature. The larvae of this species have not bee 

previously described. 

Distinguishing Features. We have identified only tM 

larvae as R. boleoides in our collections, 7.5 and 8.7 mIl

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 bod

laterally except the tail tip. A few ventral midline mela
Table 36.-Measurements (mm) of young of Radulinus asprellus. (Specimens between dashed lines are undergoing 

notochord flexion.) 

Body depth 2d preoperBody 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 1.8 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 

Rodulinus boleo/des 

Figure 28.-Larva of Radulinus boleoides. 

Table 37.-Meristics from larvae of Radulinus boleoides. (Specimens are undergoing notochord flexion.) 

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. 

Dorsal Dorsal Anal Pectoral Pelvic fin PreoperBody fin fm fin fin rays spine cular 

length spines rays rays Left Right and rays spines 

7.5 \lllI 20 15 18 N ' Buds 

8.7 IX 16 20 19 N I,3 

'Present but not countable as distinct melanophores. 

'N = Not examined. 

Ventral 

midline 

melanophores 1 

N 

N 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 depth 2d preoperBody Head Snout Eye Snout to at pectoral Pectoral cular Parietal 

length length length diameter anus length fin base fin length spine length spine length 

7.5 2.4 0.58 0.70 4.4 1.7 0.98 

8.7 2.6 0.64 0.78 5.0 2.0 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 similarity of these larvae to R. asprellus and the major differences in body proportions which were obviously not 

caused by distortion. The only other species of Radulinus 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 melanophores 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 

ventral body margins and no distinct line of melan 

phores occurs along the lateral midline. Some melan 

phores occur on the ventral finfold and occasionally 

the dorsal finfold. The preanal finfold is dark 

pigmented. Melanophores are present on the base of t 

pectoral fin. The caudal peduncle remains un pigment 

through the pelagic period. Body pigment changes litt! 

with development except as transformation approach 

pigment is added to the spinous dorsal fin and pect 

ral fin base, and pigment bands begin to show on t 

body. 

Only 1 spine develops on the dorsal margin of pr 

opercle, evident by ~ 10 mm. Smaller spiny projectio 

appear along the preopercular margin which are roo 

fied scales developing over the head region. A postocul 

spine is apparent over the eye by ~8 mm and becom 

rather pronounced in larger specimens. A parietal and 

nuchal spine develop and fuse together appearing as on 

Two spmes become obvious in the posttemporal-supr~ 

cleithral region and a cleithral spine and nasal spine 

evident on our largest specimens. 

Body shape is characteristic. Larvae are deep bodie 

increasing from 29 to 40c 0 SL between 8 and 12 m 

Snout to anus distance is >60('0 SL. By 10 mm, larv 

begin to resemble adults in form. The larvae have a di 

tinct pigmented preanal finfold which remains eviden 

anterior to the anal opening at least until -12 mm. 

our knowledge, a preanal fmfold has been reported 

Scorpaemchthys marmoratus (O'Connell 1953), rIem 

tripterus americanus (Warfel and Merriman 1944; F 

man 1976), possibly H. uillosus as indicated on a fi 

by Okiyama and Sando (1976), Myoxocephalus quadr 

cornis, M. scorpius (Khan 1972), possibly M. aenae 

based on figures by Lund and Marcy (1975), possibl 

Gymnocanthus tricuspis and G. herzensteini based 0 

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;" = Not examined.] 

Pelvic Ventral Caudal fm ra~s 

Anal Pectoral fm Preoper- midline ... Dorsal Ventral Vertebrae BranchiBody Dorsal fin fm fm ra~s 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 

8.4 -14B-** 6B - Buds N N N N N N N 

- - - - - --- ------- - ---- -------------------------- ----------------------------- ----------------------------------

10.6 VITI 13 6 16 16 1,3 N N N N N N N 

11.7 VIII 12 6 15 16 1,3 N N N N N N N N 

11.8* VIII 13 7 16 16 1,3 N 6 5 12 15 6 

Table 40.-Measurements (mm) of larvae of Rhamphocottus richardsoni. (Specimen above dashed line is undergoing 

notochord flexion.) 

Body 

length 

8.4 

10.6 

11.7 

Head 

length 

2.6 

4.1 

4.8 

Snout 

length 

0.55 

1.0 

1.0 

Eye 

diameter 

0.94 

1.1 

1.3 

Snout to 

anus length 

5.5 

7.7 

8.1 

Body depth 

at pectoral 

fin base 

46 

2.4 

4.0 

4.7 

Pectoral 

fin length 

1.0 

2.2 

2.6 

2d preopercular 

spine length 

Parietal 

spine length 

0.43 

0.35 
Rhomphocottus 

Figure ')9 _ . - Young 0 f Rhamphoco 

47 

richardson! 

8.4mm 

!!WI rtC . hardsoni. 

NL 

IO.6mm SL 

117mm SL 
phrys bison (Misitano 1978), and E. bubalis and possibly 

E. lilljeborgi (Russell 1976), within the Cottldae. 

Discussion. Rhamphocottw; is a monotypic genus. The 

larvae are distinctive . They share the character of having 

a rather heavily pigmented body, except for the tail ttp, 

with a number of other northeast Pacific species, as di -

cussed under Radulinus asprellus. but their characteristic body shape renders them outstanding except 

perhaps at the smallest sizes. The heavily pigment d 

preanal finfold, and presence of pigment on the anal finfold, base of the pectoral fin, and the ventral urface of 

the gut help distinguish small larvae. The body becomes 

covered with di tinctive spinelike scales while still in the 

pelagic habitat. The larvae are relatively rare in our collections with only seven identified specimens, all taken 

2-9 km off the Oregon coa t in March and April. 

Scorpaenichthys marmoratuH (Ay res) 

Lltf'ratur('. 

rom) of thi 

(Fi" un'~ :10. :1 1; 'I'u bh' II , 12) 

F,gg~ and young (5. r., 6.26, 10, 17, ~30, 

P cie were de cribed by 0' ,onnelJ (19. 

[h.,tlfl~Ulshlfl~ F,'rJlurf' Larvae arc between 4 an 

mm long at hatching, l\1o t (/1 (Jur pelagic pecimen 

<I:) mm. By -[:j·14 mm they have a juvenile a ppe 

ume und filii\' lormed fin . HowI'ver, we have capture 

~l.fi mm penmen peiaglcully and a 35 mm pecimen 

a nell ton tow !.J kill of! • lewport. Or 'g, 0' onnell (19 

reported p 'nmen a larg 2lJ mm were collected wi 

dip nl' , 

LlIn'a are very heavily pigmented over m t of t 

body. u wIlly appearing d!lrker than lurvae of Radulm 

Scorpoen/ch/hys mormoro/us 

- -.--~ 53mm NL 

7.5mm NL 

) 

8 .6mm NL 

Figure 30.-Larvae of Scorpaenichthys marmoratus. 

48 
Scorpaen/ch/hys marmora/us 

8.7mm SL 

IO.4mm SL 

13.8mm SL 

Figure 31.-Young of Scorpaenichthys marmoratus. 

asprellui 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 finfold, and body finfold are unpigmented. With development, melanophores are added on the head and laterally 

except along the ventral margin of the gut which remains 

49 

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 generally 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 on1y the upper 3 

spines become pronounced. They are first noticeable ~8 
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 posttemporalsupracleithral region, and the cleithral region, but spines 

never form. A nasal spine becomes obvious by ~13-

mm. 

Preflexion larvae are relatively slender with bo( 

depth at pectoral fin base ranging from 17 to 19% s: 

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.) 

Ventral Caudal fin rays 

Anal Pectoral Preoper- midline*" Dorsal Ventral Vertebrae 

Body Dorsal fin fin fin rays 

Pelvic 

fin 

spine 

& rays 

cular melano- Second- Prin- Prin- Second- Abdom- CauBranchiostegal 

length Spines Rays rays Left Right spines phores ary cipal cipal ary inal dal rays 

5.1 ' 

5.3 

5.4 

5.8 

6.1 

6.3 

6.6* 

7.5 

7.5' 

7.7* 

8.1 

8.1 

8.2' 

8.6 

8.7 

9.1 * 

9.3 

10.1' 

10.4 

10.4 

10.7 

10.9' 

11.5 

12.2' 

13.8 

13.8* 

14.7 

21.6 

-15B-'* 12B 

-16B-*' 13B 

-15-** 13 14 

12 

-16-*' 13 14 

X 18 14 13 

XI 17 12 15 

XI 15 11 14 

XI 17 12 15 

XI 17 14 15 

X 16 12 14 

XI 17 14 16 

XI 17 12 15 

XI 17 13 N 

XI 15 13 15 

X 16 13 15 

14 

10 

15 

13 

15 

14 

15 

15 

15 

16 

15 

15 

15 

15 

Buds 

Buds 

Buds 

Buds 

1,3 

1,5 

1,5 

1,5 

1,5 

1,5 

1,5 

N 

N 

N 

N 

2 

4 

N 

2 

4 

N 

4 

N 

4 

4 

4 

N 

4 

N 

4 

N 

4 

4 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

5 

N 

N 

N 

7 

N 

9 

N 

9 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

6 

N 

6 

N 

N 

N 

6 

N 

6 

N 

6 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

6 

N 

6 

N 

N 

N 

6 

N 

6 

N 

6 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

2 

N 

N 

N 

6 

N 

5 

N 

9 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

14 

N 

14 

N 

N 

N 

15 

N 

15 

N 

15 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

16 

N 

19 

N 

N 

N 

21 

N 

21 

N 

21 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

N 

6 

N 

6 

N 

N 

N 

6 

N 

6 

N 

6 

N 

N 

Table 42.-Measurements (mm) of young of Scorpaenichthys marmoratus_ (Specimens between dashed lines are 

undergoing notochord flexion.) 

Body 

length 

5.3 

5.4 

5.8 

6.1 

6.3 

Head 

length 

1.1 

1.2 

1.3 

1.4 

1.3 

Snout 

length 

0.27 

0.27 

0.26 

0.30 

0.36 

Eye Snout to 

diameter anus length 

0.44 2.8 

0.56 2.9 

0.56 3.1 

0.56 3.1 

0.56 3.1 

Body depth 2d preoperat pectoral 

fin base 

0.90 

0.98 

0.98 

1.1 

1.2 

Pectoral 

fin length 

0.44 

0.40 

0.43 

0.48 

0.46 

cular 

spine length 

Parietal 

spine length 

._--------------------- ----------- ---------------------- ------------------- -

7.5 

8.1 

8.1 

8.6 

8.7 

1.6 

1.9 

2.5 

1.9 

2.2 

0.27 

0.31 

0.39 

0.35 

0.39 

0.66 

0.82 

1.05 

0.68 

0.98 

3.7 

4.6 

4.6 

4.2 

5.2 

1.2 

1.9 

2.1 

1.6 

2.4 

0.82 

0.55 

0.82 

0.70 

0.96 

----------------------------

0.32 

0.42 

---------------------------------------------- --- ---- ----------------------------------------------------

9.3 

10.4 

10.4 

10.7 

11.5 

13.8 

14'.7 

21.6 

3.1 

3.8 

3.1 

3.4 

3.8 

5.4 

5.4 

8.8 

0.40 

0.72 

0.48 

0.48 

0.40 

0.96 

0.88 

1.92 

1.4 

2.2 

1.4 

1.7 

1.8 

2.2 

2.5 

2.7 

5.8 

6.2 

5.9 

6.5 

7.4 

8.9 

9.7 

15.5 

50 

3.2 

3.3 

3.3 

3.8 

3.8 

4.6 

5.2 

8.2 

1.6 

1.8 

1.8 

2.0 

2.2 

3.0 

3.4 

6.2 

0.70 

0.90 

0.78 

0.78 

1.01 

1.24 

1.36 

1.44 

Bump 

Bump 
ey become much deeper bodied with development 

'th a body depth of 32-35% SL in postflexion stages. 

out to anus length is relatively long, increasing from 

-53% SL in preflexion larvae to 57-66% SL in postexion larvae. The snout develops a rounded appearce, never pointed as in Radulinus sp. An unpigmented 

eanal finfold remains evident up to ~13-14 mm. The 

esence of such a preanal finfold has been reported for 

latively few cottid genera (see Discussion under Rhamhocottus richardsoni). 

iscussion. Scorpaenichthys is a monotypic genus. The 

ae are rather distinctive and one of the most heavily 

[gmented forms of northeast Pacific cottid species. 

all larvae appear somewhat similar to Radulinus spp. 

t have lateral pigment extending further toward the 

Iii tip, no ventral midline melanophores posterior to the 

teral pigment and usually no distinct line of melanohores along the lateral midline. Occasionally newly 

atched larvae will have very few melanophores on the 

teral body surface causing them to resemble small 

remilepidotus spinosus. However, their gut is longer and 

e dorsal and ventral midline pigment does not extend 

s far onto the tail tip as in H. spinosus. 

Larvae of S. marmoratus appear to be strongly assoated with the neuston. O'Connell (1953) reported dip 

t collections contained specimens 7.4-29 mm long, and 

e have taken specimens 4.0-35 mm in neuston tows off 

regon. One 10-min neuston tow taken 9 km off Newrt, Oreg., ill March contained 829 larvae, mostly 4-5 

m long and recently hatched. Richardson and Pearcy 

L977) reported them to be a coastal form, as the larvae 

ere only taken within 28 km of the coast in oblique 

lankton tows. However, we have since collected them in 

he neuston as far as 65 km offshore, well beyond the 

ntinental shelf-slope break. 

Cottoid Type A 

(Figure 32; Tables 43, 44) 

iterature. No published descriptions of this form exist 

lthough illustrations were given by Richardson and 

ond9 as "Globby Type A." These larvae were called 

'Richardson. S. L .• and C. E. Bond. 1978. Two unusual cottoid 

ishes from the northeast Pacific. Unpubl. manuscr.. 6 p. Gulf Coast 

search Laboratory. Ocean Springs, MS 39564, or Department of Fishries and ~Vildlife. Oregon State University. Corvallis, OR 97331. 

Psychrolutes-like sp. 1 by Richard on (footnote 4) and 

Richardson and Pearcy (1977). 

Distinguishing Features. Three larval pecimen (9. , 

12.8, 13.4 mm) of this unusual looking form have been 

collected off Oregon. The smallest pecimen i undergoing notochord flexion and the two larger specimens are 

postflexion larvae. The largest larva, which was tained, 

has the full complement of dorsal, anal, pectoral, and 

pelvic fin rays but secondary caudal rays are till 

developing and the vertebral centra are not completely 

ossified, thus transformation occurs at a larger lze. 

The larvae are pigmented over the entire body except 

at the tail tip. The basal portion of the pectoral fin i 

pigmented, but its distal portion and the dorsal and anal 

finfolds are unpigmented. 

No spines are apparent in the head region. The belly i 

covered with small, thumb-tack prickles which extend 

dorsolaterally 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. Thi condition may be variable as reported in adults in a Southern 

Hemisphere cottoid genus N eophrynichthys ( elson 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 fm rays and the possession of seven branchlostegal 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 pflckles in 

the belly region similar to these larvae. Based on thi 

information alone our Cottoid Type A larvae would appear 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 po e ion of 

belly prickles has lead to uncertainty regarding it identity. 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 Ill'. ion.) 

[* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable: D = Damagl'd.. =. 'ot I'xam.ned ) 

Pelvic Ventral Caudal fin ra\ 

Anal Pl'ctoral fin Preoper· midline Dorsal Ventral \ 'enehrae Branch.· 

Body Dorsal fin fin fin rays spine cular Second· Prin· Prin, econd· Abdom· legal 

length Spines Rays rays Left Right & rays ra} 

9.8 ·15·" 11 23 23 

12.8 ·16·" 11 24 D ~ :\ :-.; 

~3.~4_· ___ \1ll ~ __ ~~~O __ ~1~4 __ ~26~~26~ __ -2I.3~ ____________________ ~6 i 4 12 

51 
Cottoid Type A 

/.."r ----"""--- -, - ... _ 

.' ~ 

. ---

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 notochord 

flexion.) 

Body depth 2d preoperBody Head Snout Eye Snout to at pectoral Pectoral cular Parietal 

length length length diameter anus length fm base fin length spine length spine length 

9.8 3.6 0.66 1.2 5.7 4.1 1.7 

12.8 6.1 1.5 1.7 10.1 7.2 3.7 

52 
dy depth (51 % and 31 % SL, respectively) and snout to 

us length (61% and 51% SL, respectively). The 

lidentified 26.4 mm juvenile agrees much more closely 

th our Cottoid Type A larvae than with the 35 mm P. 

rictus juvenile. It seems unlikely that a drastic change 

body proportion would occur during the juvenile 

.riod between 26 and 35 mm. And the 26.4 mm specin does not appear to be distorted, based on radiophs. It also seems unlikely that two such unusual 

rms would exist in the northeast Pacific that agree so 

ell in meristics and belly prickles. However, the identiation of our Cottoid Type A larvae as P. phrictus can-

~t be positively made until additional material becomes 

railable to provide a conclusive linkage between our larle and identified juveniles. 

If our Cottoid Type A larvae are found to be P. 

rictus, then placement of this species in the genus Psyrolutes is obviously in error based on the extreme dismilarity of these larvae and those of P. paradox us de-

:ribed by Blackburn (1973) (10.3 mm SL) and Marliave 

.975) (10.5, 13, 14 mm TL), and its generic position 

10uld be reexamined. Alternatively, these larvae could 

lpresent a new species and possibly a new genus of cotid fish from the northeast Pacific. The term "cottoid" 

as been used here because of the dissimilarity of these 

lrvae to any known larvae in the Family Cottidae. 

NOTES ON OTHER 

NORTHEAST PACIFIC COTTID LARVAE 

Larvae are known for 14 additional species of northst 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, ?Malacocottus, 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 fms are unpigmented 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 beginning 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. marmoratus 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. marmoratus. The largest specimen reported from plankton 

collections is 25.5 mm TL and is apparently transformed, or nearly so. 

Table 45.-Additional northeast Pacific coUid species for which larvae-and young are known. 

Species Reference Size illustrated --------------~------------~~~==~---

Artedius lateralis 

Blepsias cirrhosus 

Clinocottus analis 

Clinocottus recaluus 

Dasycottus setiger 

Gilbertidia sigalutes 

H emilepidotus jordani 

Hemilepidotus zapus 

?Malacocottus zonurus 

-Type 1 

Myoxocephalus polyacanthocephalus 

Oligocottus maculosus 

o ligocottus snyderi 

Orthonopias triacis 

Psychrolutes paradoxus 

1 See text footnote 9. 

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 19781 

Richardson (unpubl. data) 

Blackburn 1973 

Stein 1972 

Stein 1973 

Stein 1972 

Bolin 1941 

Blackburn 1973 

Marliave 1975 

53 

4.1 mm SL 

4,8, 11, 14 mm TL 

12.2mm SL 

10, 14, 19 25.5 mm TL 

-4mm 

-4mm 

4.6,5.0,7.6,8.3,9.9,10.8,18.0, 

24.3 mmTL 

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 

mmSL 

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 

-3-4mmSL 

10.3 mm SL 

10.5, 13, 14, 13 mm TL 
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 appearance, 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 pectoral 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 presumed 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 Cilbertidia sigalutes and Psychrolutes 

paradox us 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. Postanal 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. paradox us than C. 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 2: 19 for P. paradoxus and ~18 for C. sigalutes. These are usually countable on all but the smallest larvae. Additional definitive 

differences between the smallest larvae of these two species, 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. paradox us have more melanin in the pectoral 

fins which obscures their orange color while the pectoral 

fins of C. 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 variation of those patterns remain to be defined. Pectoral fin 

pigment differences are apparent in larger preserved 

specimens with those of C. sigalutes being much darker 

and having unpigmented circles near the margin. 

Marliave also stated that P. paradox us larvae develop a 

notch dorsally in the caudal finfold during fin ray formation whereas C. sigalutes larvae do not. This was not obvious on all specimens we have examined. Marliave 

observed that P. paradoxus settles from the plankton 

-13-14 mm TL while C. sigalutes remains pelagic until 

>20 mm TL. Larvae of an additional form from the Gulf 

of Alaska and Bering Sea, tentatively called Malacocottus zonurus - Type 1 by Richardson and Bond (footnote 9), are also very similar in appearance to larvae of 

G. sigalutes and P. paradox us. Small larvae of all three 

are nearly identical but larvae of M. zonurus - Type 1 

develop 4 preopercular spines with an anteriorly directed accessory spine at the base of the second spine. 

This secondary spine is visible on larvae by ~7 mm upon 

fl4 

close examination. Pectoral fin ray counts, ~20-22, wi 

further separate it from C. sigalutes. Often the pelvic fi 

of M. zonurus - Type 1 appear to be withdrawn int 

pockets in the skin, although this condition is variabl 

This form has very loose skin and remains pelagic at lea 

until ~24 mm. Positive identification of these larvae 

M . zonurus awaits resolution of apparent taxonomi 

problems in the genus Malacocottus (Howe and Ric 

ardson footnote 3; Richardson and Bond footnote 9). 

In larvae of Myoxocephalus polyacanthocephal 

which hatch ~7 mm, melanophores cover most of t 

anterior one-half to two-thirds of the body except t 

ventral surface of the abdomen. The posterior one-thi 

of the body remains conspicuously unpigmented excep 

for a series of small melanophores along the ventr~ 

midline which extends to near the tail tip. The pigme 

pattern is similar to other dark cottid larvae, e.g., Rad 

linus asprellus, Rhamphocottus richardson 

Scorpaenichthys marmoratus, Blepsias cirrhosus, exce 

that a greater proportion of the tail region remains un 

pigmented laterally and the unpigmented tail region al 

retains a series of ventral midline melanophores. Til 

head develops prominent spines including at least 4 pr 

opercular, a parietal, a nuchal, a postocular, and severa 

in the posttemporal-supracleithral region. 

Early larvae of Orthonopias triacis resemble those 

Artedius spp. with which it is closely related (Howe an 

Richardson footnote 3). At hatching they have pigme 

over the gut and a series of ~35 post anal ventral midli 

melanophores. Soon after hatching pigment is added 

the interorbital region and on the nape. Apparently n 

gut diverticula are present. Characters given in the lite 

ature are inadequate to distinguish them from Artedi 

harringtoni except for possibly a higher number of ven 

tral midline melanophores. 

ACKNOWLEDGMENTS 

Examination of larval cottids from other areas of th 

northeast Pacific was helpful in working out identifica 

tions of our Oregon cottid larvae. We thank the followin 

people with collections from particular areas: north en 

California-David W. Rice (Lawrence Livermore Labo 

ratory); Washington and Puget Sound-Jean R. Du 

and Kenneth D. Waldron ( orthwest and Alaska Fish 

eries Center, National Marine Fisheries Service, NOAA

Lawrence Moulton (University of Washington), Jame

Blackburn (Alaska Department of Fish and Gam 

formerly of University of Washington), Andrew Lam~ 

(Pacific Environment Institute, Vancouver), British Co

umbia-W. E. Barraclough (Fisheries Research Board 0 

Canada, Nanaimo) and Daniel J. Faber (Canadi 

Oceanographic Identification Center); Bering Sea an

Gulf of Alaska-Pat Wagner and George Mueller (Un

versity of Alaska). Jean R. Dunn provided reared larva

of Hemilepidotus hemilepidotus. Kevin Howe, Orego

State University, provided transforming and juvenil

cottids collected in Oregon tidepools and general infor

mation on tidepool collections. James Blackburn an

Jeffrey B. Marliave, Vancouver Public Aquarium, pro
ded specimens of Gilbertidia sigalutes and Psychrotes paradox us for comparative examination. Many 

eople helped in one way or another in working out larval 

ottid identification problems particularly E. H. Ahl-

,rom, NOAA, NMFS Southwest Fisheries Center; 

mes Blackburn; Carl E. Bond, Oregon State Uniersity; Jean R. Dunn; Kevin Howe; Joanne L. Laroche, 

regon State University; Wayne L. Laroche, Oregon 

tate University; Kenneth Waldron; and Sharon Roe, 

Drmerly of Oregon State University. We thank E. H. 

lstrom and Carl E. Bond for reviewing the manucript and offering h,elpful comments. Special thanks are 

octended to Jean R. Dunn whose efforts made this paper 

ossible. This research was supported in part by NOAAMFS Contract No. 03-78-M02-120, 1 January 1978 to 

o September 1978. 

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on. parate heet from the text For 

pnJlCeIl8d material live author, year title 

tDI_"'~ DUmber of ..... ' and where It filed-agen 

lte name In text and footnote 

...... : John J Jon hery BIOI0(p8t, nppe lnatl 

iGlrtfo._apapl!ty, La Jolla, 92037, pen common 21