FS ajr/fc f<,?o 430 s Utk o* " NOAA Technical Report cular 430 Guide to Identification of Some Sculpin (Cottidae) Larvae from Marine and Brackish Waters off Oregon and Adjacent Areas in The Northeast Pacific Sally L. Richardson and Betsy B. Washington January 1980 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels for optimum use of the resources. 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For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, DC 20402; Stock No. 003-017-00412-1. 407. Distribution of decapod Crustacea off northeastern United States based on specimens at the Northeast Fisheries Center, Woods Hole, Massachusetts. By Austin B. Williams and Roland L. Wigley. De- cember 1977, iii + 44 p., 2 figs., 1 table, 57 charts. 408. Collection of tuna baitfish papers. (20 papers.) By Richard S. Shomura (editor). December 1977, iii + 167 p. 409. Marine flora and fauna of the northeastern United States. Copepoda: Cyclopoids parasitic on fishes. By Ju-Shey Ho. February 1978, iii + 12 p., 17 figs. 410. The 1976 Ceratium tripos bloom in the New York Bight: Causes and consequences. By Thomas C. Malone. May 1978, iv + 14 p., 17 figs., 1 table. 411. Systematics and biology of the tilefishes ( p erciformes: Branchio- stegidae and Malacanthidae), wuh descriptions of two new species. By James K. Dooley. April 1978, v + 78 p., 4^ figs., 26 tables. 412. Synopsis of biological data on the red porgy, Pagrus pagrus (Lin- naeus). By Charles S. Manooch III and William W. Hassler. May 1978, iii + 19 p., 12 figs., 7 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, DC 20402; Stock No. 003-017-00418-0. 413. Marine flora and fauna of the northeastern United States. Crustacea: Branchiura. By Roger F. Cressey. May 1978, iii + 10 p., 15 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, DC 20402; Stock No. 003-017-004.1-8. 414. Synopsis of biological data for the winter flounder. Pseudopleuro- nectes americanus (Walbaun.;. By Grace Klein-MacPhee. November 1978, iii + 43 p., 21 figs., 28 tables. 403. Marine flora and fauna of the northeastern United States. Sipun- cula. By Edward B. Cutler. July 1977, iii + 7 p., 6 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, DC 20402; Stock No. 003-012-00404-0. 404. Revision of the sea basses of the genus Diplectrum (Pisces: Ser- ranidae). By Stephen A. Bortone. September 1977, v + 49 p., 15 figs., 9 tables. 405. Marine flora and fauna of the northeastern United States. Echino- dermata: Holothuroidea. By David L. Pawson. September 1977, iii + 15 p., 3 figs. For sale by the Superintendent of Documents, U.S. Gov- 415. A basis for classifying western Atlantic Sciaenidae (Teleostei: Perciformes). By Labbish Ning Chao. September 1978, v + 64 p., 41 figs., 1 table. 416. Ocean variability: Effects on U.S. marine fishery resources - 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 Al MOSp, NT Of C Guide to Identification of Some Sculpin (Cottidae) Larvae from Marine and Brackish Waters off Oregon and Adjacent Areas in the Northeast Pacific Sally L. Richardson and Betsy B. Washington January 1980 2? (X o U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps, Secretary National Oceanic and Atmospheric Administration Richard A. Frank, Administrator National Marine Fisheries Service Terry L. Leitzell, Assistant Administrator for Fisheries For Sale by the Superintendent of Documents, U.S. Government Printing Office Washington, DC. 20402 - Stock No. 003-017-00472-4 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. 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CONTENTS Introduction 1 Methods 2 Useful characters 3 Order of presentation • 4 Artedius harringtoni (Starks) 4 Artedius Type 2 8 Clinocottus acuticeps (Gilbert) 12 Cottidae Type 1 15 Cottidae Type 2 IV Cottidae Type 3 18 Trigolops sp 20 Chitonotus pugetensis (Steindachner) 22 Cottus asper Richardson 24 Enophrys bison (Girard) 25 Icelinus spp 27 Leptocottus armatus Girard 31 Paricelinus hopliticus Eigenmann and Eigenmann 32 Hemilepidotus hemilepidotus (Tilesius) 34 Hemilepidotus spinosus (Ayres) 37 Nautichthys oculofasciatus (Girard) 40 Radulinus asprellus Gilbert 42 Radulinus boleoides Gilbert 44 Rhamphocottus richardsoni Giinther 46 Scorpaenichthys marmoratus (Ayres) 48 Cottoid Type A 51 Notes on other northeast Pacific cottid larvae 53 Acknowledgments 54 Literature cited 55 Figures 1. Larvae of Artedius harringtoni 6 2. Larvae of Artedius harringtoni 7 3. Larvae of Artedius Type 2. The second figure from the top is a dorsal view showing gut diverticula ... 9 4. Larvae of Artedius Type 2 10 5. Larvae of Clinocottus acuticeps. The second figure from the top is a ventral view showing hindgut diverticula 12 6. Young of Clinocottus acuticeps 13 7. Larvae of Cottidae Type 1 16 8. Larvae of Cottidae Type 2 17 9. Larvae of Cottidae Type 3 19 10. Larvae of Triglops sp 21 11. Larvae of Chitonotus pugetensis 22 12. Young of Chitonotus pugetensis 23 13. Larvae of Cottus asper 25 14. Young of Enophrys bison 26 15. Larvae of Icelinus spp 28 16. Young of Icelinus spp 29 17. Young of Icelinus spp 30 18. Larvae of Leptocottus armatus 31 19. Larvae of Paricelinus hopliticus 33 20. Young of Paricelinus hopliticus 34 21. Larvae of Hemilepidotus hemilepidotus 35 22. Young of Hemilepidotus hemilepidotus 36 23. Larvae of Hemilepidotus spinosus 38 24. Young of Hemilepidotus spinosus 39 25. Larvae of Nautichthys oculofasciatus 41 26. Larvae of Radulinus asprellus 43 27. Young of Radulinus asprellus 44 iii 28. Larva of Radulinus boleoides 45 29. Young of Rhamphocottus richardsoni 47 30. Larvae of Scorpaenichthys marmoratus 48 31. Young of Scorpaenichthys marmoratus 49 32. Larvae of Cottoid Type A 52 Tables 1. Species of adult cottids whose range of distribution reportedly includes marine or brackish waters off Oregon and their meristics 2 2. Groupings of cottid larvae described in this guide based on preopercular spines and pigment pat- terns 5 3. Meristics from larvae of Artedius harringtoni 5 4. Measurements (mm) of larvae of Artedius harringtoni 5 5. Meristics from larvae of Artedius Type 2 8 6. Measurements (mm) of larvae of Artedius Type 2 8 7. Meristics from young of Clinocottus acuticeps 14 8. Measurements (mm) of larvae of Clinocottus acuticeps 14 9. Meristics of larvae of Cottidae Type 1 15 10. Measurements (mm) of larvae of Cottidae Type 1 , 15 11. Meristics from larvae of Cottidae Type 2 18 12. Measurements (mm) of larvae of Cottidae Type 2 18 13. Meristics from larvae of Cottidae Type 3 20 14. Measurements (mm) of larvae of Cottidae Type 3 20 15. Meristics from larvae of Triglops sp 20 16. Measurements (mm) of larvae of Triglops sp 20 17. Meristics from larvae of Chitonotus pugetensis 23 18. Measurements (mm) of larvae of Chitonotus pugetensis 24 19. Meristics from larvae of Cottus asper 24 20. Measurements (mm) of larvae of Cottus asper 24 21. Meristics from larvae of Enophrys bison 25 22. Measurements (mm) of larvae of Enophrys bison 27 23. Meristics from larvae of Icelinus spp 30 24. Measurements (mm) of larvae of Icelinus spp 31 25. Meristics from larvae of Leptocottus armatus 32 26. Measurements (mm) of larvae of Leptocottus armatus 32 27. Meristics from young of Paricelinus hopliticus 32 28. Measurements (mm) of young of Paricelinus hopliticus 33 29. Meristics from young of Hemilepidotus hemilepidotus 34 30. Measurements (mm) of young Hemilepidotus hemilepidotus 36 31. Meristics from young of Hemilepidotus spinosus 40 32. Measurements (mm) of young of Hemilepidotus spinosus 41 33. Meristics from larvae of Nautichthys oculofasciatus 42 34. Measurements (mm) of larvae of Nautichthys oculofasciatus 42 35. Meristics from young of Radulinus asprellus 42 36. Measurements (mm) of young of Radulinus asprellus 45 37. Meristics from larvae of Radulinus boleoides 45 38. Measurements (mm) of larvae of Radulinus boleoides 45 39. Meristics from larvae of Rhamphocottus richardsoni 46 40. Measurements (mm) of larvae of Rhamphocottus richardoni 46 41. Meristics from young of Scorpaenichthys marmoratus 50 42. Measurements (mm) of young of Scorpaenichthys marmoratus 50 43. Meristics from larvae of Cottoid Type A 51 44. Measurements (mm) of larvae of Cottoid Type A 52 45. Additional northeast Pacific cottid species for which larvae are known 53 IV Guide to Identification of Some Sculpin (Cottidae) Larvae from Marine and Brackish Waters off Oregon and Adjacent Areas in the Northeast Pacific Sally L. Richardson 1 and Betsy B. Washington 2 ABSTRACT Knowledge of identification of cottid larvae taken in plankton collections off Oregon is sum- marized. Included are descriptions of 21 kinds of cottid larvae representing at least 14 of the 40 genera of cottids reported to occur in the northeast Pacific between Baja California and the Aleutian Islands. Larvae of Artedius harringtoni, Chitonotus pugetensis, Clinocottus acuticeps, Icelinus spp., Paricelinus hopliticus, Radulinus asprellus, R. boleoides, and four unidentified types are described for the first time. A larval series of a second kind of Artedius is described with a discusion of problems of specific identification. New and comparative information is provided for larvae of Cottus asper, Enophrys bison, Hemilepidotus hemilepidotus, H. spinosus, Leptocottus armatus, Nautiehthys oeulo- fasciatus, Rhamphocottus richardsoni, Scorpaenichthys marmoratus, and Triglops sp. Each descriptive account contains illustrations, a literature summary, distinguishing features (including size at hatching and transformation, pigmentation, head spination, general morphology), and a discussion of identification problems. Characters found to be most useful in distinguishing cot- tid larvae include morphology, pigmentation, head spination, and meristics. Literature summaries are also provided for northeast Pacific cottid larvae in 8 additional genera: Blepsias, Dasycottus, Gilbertidia, ?Malacoeottus, Myoxocephalus, Oligocottus, Orthonopias, Psychrolutes. INTRODUCTION The cottids are one of the most speciose and diverse groups of fishes inhabiting coastal waters of the north- east Pacific Ocean with 90 species in 40 genera reported to occur from Baja California to the Aleutian Islands (Howe and Richardson 1 ). The range of 36 of these species in 20 genera reportedly includes waters off Oregon (Table 1). Adult cottids range in size from the 5 cm Artedius meanyi, Clinocottus acuticeps, and Oligocottus rimensis to the 76 cm Myoxocephalus poly- acanthoceihalus and Scorpaenichthys marmoratus (Hart 1973). Larger forms in the genera Gymno- canthus, Hemilepidotus, Hemitripterus, Melletes, and Myoxocephalus are fished commercially by the U.S.S.R. and Japan (Gorbonova 1964; Fedorov 1973). Off Oregon several species are taken frequently by sports fishermen, e.g., Enophrys bison, Hemilepidotus hemilepidotus, H. spinosus, Leptocottus armatus, Scor- paenichthys marmoratus (Beardsley and Bond 1970). 'School of Oceanography, Oregon State University, Corvallis, Oreg.; present address: Gulf Coast Research Laboratory, East Beach Drive, Ocean Springs, MS 39564. 'School of Oceanography, Oregon State University, Corvallis, OR 97330. Howe, K., and S. L. Richardson. 1978. Taxonomic review and meristic variation in marine sculpins (Osteichthys: Cottidae) of the northeast Pacific Ocean. Final Report, NOAA NMFS Contract No. 03- 78-M02-120, 1 January 1978 to 30 September 1978, 142 p. Northwest and Alaska Fisheries Center, National Marine Fisheries Service, NOAA, 2725 Montlake Blvd. East, Seattle, WA 98102. Most, if not all cottid larvae hatch from demersal eggs ~(about)l-4 mm in diameter (Breder and Rosen 1966) and range from approximately 3 to 12 mm at hatching. They are usually pelagic and some are primarily neustonic, i.e., occurring at the surface of the water column, such as Scorpaenichthys marmoratus and Hemilepidotus spinosus (unpubl. data). Most cottid lar- vae probably transform to juveniles by ~20 mm SL. Cottid larvae are collected frequently in the coastal zone of temperate and boreal areas of the northeast Pacific but they are usually not abundant. In a l'/a yr survey off Oregon they occurred in 80' 7c of the samples but comprised only 8 c h of the fish larvae captured in a coastal assemblage of larval fishes 2-28 km offshore (Richardson 1977 4 ; Richardson and Pearcy 1977). They were taken in all months of the year in that study. A wide variety of larval forms is demonstrated within the cottid family. Identification of the larvae is often dif- ficult because of the large number of species in the fami- ly, because of difficulty in establishing developmental series from the small numbers of larvae in plankton samples, and because relatively few species have been described in detail. This report summarizes knowledge of identification of cottid larvae taken primarily in plankton and neuston ^Richardson, S. L. 1977. Larval fishes in ocean waters off Yaquina Bay, Oregon: abundance, distribution, and seasonality January 1971 to August 1972. Oreg. State Univ., Sea Grant Coll. Prog. Publ. No. ORESU-T-77-003, 73 p. Table 1. — Species of adult cottids whose range of distribution reportedly includes marine or brackish waters off Oregon and their meristics. 1 Branchiostegal rays num- ber 6 unless otherwise noted. Dorsal Dorsal fin Anal fin soft fin Pectoral Pelvic Total Species spines 2 rays 2 rays- fin rays fin rays vertebrae Artedius eorallinus VIII-IX 15-16 12-13 14-16 3 31-33 Artedius fenestralis VIII -IX 16-18 12-14 14-16 3 32-35 Artedius harringtoni' IX 15-18 10-14 13-15 3 32-34 Artedius lateralis VII-X 15-17 12-14 14-16 3 32-34 Artedius meanyi IX-X 14-17 10-12 14-16 2-3* 33-35 Artedius notospilotus IX-X 14-16 11-13 14-17 3 32-34 Ascelichthys rhodorus VII-X 17-19 13-16 16-18 33-36 Blepsias cirrhosus VI-VIII 20-24 18-21 11-13 3 37-38 Chitonotus pugetensis VIII- XI 14-17 14-17 16-18 2-3* 35-36 Clinocottus acuticeps VII-IX 13-17 9-13 13-15 3 31-33 Clinocottus embryum vni-x 14-17 9-12 12-15 3 33-34 Clinocottus globiceps vm-x 13-17 11-12 13-15 3 32-34 Clinocottus recaluus VIII-IX 14-16 9-13 13-15 3 32-33 Cottus aleuticus VIII-X 16-20 12-16 13-16 4 34-39 Cottus asper" VII-XI 18-21 14-18 14-17 4 34-39 Enophrys bison VII-IX 9-13 8-10 15-17 3-4* 29-31 Hemilepidotus hemilepidotus X-XIII 18-20 13-16 15-17 4 35-37 Hemilepidotus spinosus XXI 18-20 14-16 14-16 4 35-37 Icelinus burchami XXI 16-18 12-14 16-19 2 35-37 Icelinus filamentosus X-XII 15-17 13-16 16-18 2 34-37 Icelinus oculatus XI 15-17 13-14 17 2 37 Icelinus tenuis IX-XI 16-18 14-17 15-17 2 37-39 Jordania zonope XVII-XVIII 15-18 22-24 13-15 4-5* 46-48 Leptocottus armatus VI-VIII 15-20 15-20 17-20 4 35-39 Nautichthys oculofasciatus VIII-IX 27-30 16-21 13-14 3 40-41 Oligocottus maculosus VIII-IX 15-18 12-14 12-15 3 33-34 Oligocottus rimensis VIII-X 16-19 13-15 13-15 3 34-37 Oligocottus snyderi VII-IX 17-20 12-15 12-15 3 34-37 Paricelinus hopliticus xii-xm 19-20 23-24 14-15 5 42 Psychrolutes phrictus 4 VII-IX 19-20 12-14 22-26 3 33-35 Radulinus asprellus VIII- X 20-23 21-25 17-20 3 38-39 Radulinus boleoides VIII-XI 20-22 21-23 18-20 3 39-40 Rhamphocottus richardsoni VII-IX 12-14 6-8 14-16 3*-4 26-28 Scorpaenichthys marmoratus VIII- XII 15-19 11-14 14-16 4-5* 35-37 Synchirus gilli VIII-X 19-21 18-21 21-24 3 38-39 Zesticelus profundorum Y VII 10-13 8-11 19-21 2*-3 25-26 'From compilation by Howe and Richardson 1978 (see text footnote 3) which incorporated data from literature sources and original counts. "Includes all fin ray elements whether or not they arise from the same pterigiophore. 'Includes hypural. 'Has 7 branchiostegal rays instead of the usual 6. 'May have 6 or 7 branchiostegal rays. *Most frequent count. collections off the Oregon coast. Included are descrip- tions and illustrations of complete or partial developmental series of 21 kinds of cottid larvae. These represent all but one form of cottid larvae from Oregon waters for which identified (as species or type) material is available in the Oregon State University Larval Fish Reference Collection. Although keys are not provided, larvae are grouped together on the basis of common lar- val characters. Occurrences, including seasonality, in- shore-offshore distribution and relative abundance of most of these cottid larvae off Oregon have been reported by Richardson (see footnote 4) and Richardson and Pearcy 1977. However, relevant new information on occurrences is presented where appropriate. Knowledge of other cottid larvae from the northeast Pacific is sum- marized from the literature and personal observation. This paper provides a basis for comparison with ad- ditional cottid species whose larvae remain to be iden- tified and described. It is primarily descriptive in nature. Examination of relationships withinin the northeast Pacific Cottidae based on larval morphology is presented elsewhere (Richardson in press). METHODS Specimens came primarily from over 1,000 plankton and neuston samples collected off Oregon since 1969, which contained over 2,000 cottid larvae. During the course of working out identifications additional mate- rial was examined from plankton collections taken off northern California, Washington, Puget Sound, British Columbia, and Alaska (see Acknowledgments). Some transforming and juvenile specimens collected from Oregon tidepools were also examined. All material had been preserved in either 5 or 10% Formalin and some specimens had been transferred to 36 or 40% isopropyl alcohol. Developmental terminology generally follows Ahlstrom et al. (1976) where the larval period is separated into three stages, preflexion, flexion, and pcstflexion, on the basis of the relative position of the notochord tip. Some subjectivity is involved in deciding the exact size at which the notochord begins to flex and when it is in a fully flexed position, but we tried to be consistent in judgment in all of the series. The transi- tion from postflexion larva to juvenile could not be precisely defined in most series because of lack of specimens in the critical size range. Generally, the transformation to juvenile is accompanied by an in- crease in body pigmentation, scale formation, and com- pletion of development of all fin rays. We refer to fish as postflexion larvae until they have attained an adult body form which characterizes them as juveniles. Counts of various body parts were usually made on unstained specimens. When adequate material was available, a developmental series or several specimens were stained using the method of Taylor (1967). In un- stained specimens fin rays were counted if any elements were visible. In stained specimens, fin rays were counted if any part absorbed stain. Fin ray counts include all ray elements whether or not they arise from the same pterygiophore. Vertebral counts include the urostyle. Measurements were made following definitions given by Richardson and Laroche (1979) except: Body length = snout tip to notochord tip preceding development of caudal fin [notochord length (NL)], then to posterior margin of hypural plate [standard length (SL)]. This applies to all lengths given in this paper unless noted otherwise. Snout length = horizontal distance from tip of snout to anterior margin of pigmented region of left eye. Eye diameter = greatest diameter of pigmented portion of eye. Preopercular spine length (second spine) = from tip to basal insertion of core of spine. Head length is abbreviated as HL. Head spine terminology in cottid larvae has not been well defined. Many spines present during larval development are not present or obvious in adults and names used for spines in adults are often inconsistent. A standard terminology for spines in larval cottids, based on studies of developmental osteology of many species, remains to be proposed. In this paper, names used have been generally adapted from those that Richardson and Laroche (1979) diagramed for larvae and juveniles of Sebastes spp. for which similar problems exist. Some deviations from their terminology for Sebastes spp. exist in cottids. Preopercular spines along the posterior margin frequently number 4 but sometime may be mul- tiple (over 5) serrations or small spines, rather than 5 as in Sebastes spp. Additional terms used here include: spines in the parietal region (several spines in the vicinity of the parietal spines of Sebastes spp., sometimes called occipital spines in adult cottids); spines in the posttemporal-supracleithral region [several spines in the vicinity of the posttemporal and supracleithral spines of Sebastes spp., the basal origin of which could not be determined on unstained cottid lar- vae (not to be confused with spiny scales that develop in this region along the lateral line)); anterior parietal spine or bump (apparently on the parietal ridge ante- rior to the principle parietal spine); midopercular spine (near the middle of the posterior margin of the opercle). Illustrations were made with the aid of camera lucida. They are intended to accurately represent numbers of myomeres, fin rays including ray elements and bases, head spines, scales, and ventral midline melanophores. Identifications in most cases could be made using adult characters, primarily meristic (Table 1; Howe and Richardson footnote 3) on the largest larvae in a series linked together by pigment and head spine patterns. Additional adult characters used to confirm identifica- tions included deep emargination between dorsal fin spines 3 and 4 in Chitonotus pugetensis, attachment of inner pelvic ray to belly by membrane in Clinocottus aeutieeps, number of dorsal scale rows and scales above the lateral line in Hemilepidotus hemilepidotus and H. spinosus, elongated first dorsal fin in Nautichthys oculo- fasciatus, and unique shape in Rhamphocottus richard- soni. Identification of larvae of Radulinus boleoides was based on their similarity to R. asprellus. Newly hatched, reared larvae of Hemilepidotus hemilepidotus, Dunn helped confirm the identification of small larvae in that series. The descriptive format is designed to present a con- cise account of a combination of characters which, together with meristics (not included in the descrip- tions), will help to distinguish each of the kinds of lar- vae. Available literature is summarized for each larval type and lengths are given for each specimen illustrated. Distinguishing features in each account include infor- mation on size at hatching and transformation, pig- mentation, head spination, and general morphology. Discussions for each kind of larvae may include taxo- nomic problems, comparisons with similar species, and new occurrence data. USEFUL CHARACTERS Characters most useful to distinguish cottid larvae include general shape of the body and snout, unusual diverticula of the gut cavity, melanistic pigment pat- terns, head spination, and meristics. Body shape can range from short and stubby (Artedius harringtoni, Clinocottus aeutieeps) to moderately long and slender (Icelinus spp., Triglops sp.). Cottid larvae have a distinc- tively coiled gut and the hindgut may trail below the body (Artedius harringtoni, Clinocottus aeutieeps). Snout to anus length is usually ~ 40-50% SL except in small (<5 mm) larvae where it may be less. However, some species have a rather long gut, >60% SL (Clinocot- 'Dunn, J. R. 1973. Unpubl. data. Northwest and Alaska Fisheries Center, National Marine Fisheries Service, NOAA, 2725 Montlake Blvd. East, Seattle, WA 98102. tus acuticeps, Rhamphocottus richardsoni). Small (<6 mm) cottid larvae have a very characteristically pro- truding snout which, with development, can vary con- siderably in its degree of pointedness (e.g., compare Scorpaenichthys marmoratus and Radulinus asprellus). Several forms have unusual gut diverticula or protru- sions from the abdominal wall {Artedius Type 2, Clinocottus acuticeps) which serve as striking characters. Pigment patterns can range from darkly pigmented forms such as Scorpaenichthys marmoratus and Radulinus asprellus where melanophores cover the entire body except the tail tip to relatively unpigmented forms such as Triglops sp. with melanophores only on the head and gut. In many species (Artedius harringtoni, Chitonotus pugetensis, Paricelinus hopliticus) the number, extent, spacing, and shape of the postanal ventral midline melanophores serve as diagnostic aids, even though these may decrease in number with develop- ment. Ranges and frequency of counts of these melano- phores within developmental periods are useful. The presence or absence of nape pigment and/or head pig- ment, particularly in small (<5 mm) larvae, may be helpful in some cases {Artedius spp., Clinocottus spp., Enophrys spp.). The pattern and extent of pigment over the gut is useful in otherwise lightly pigmented forms. It may range from relatively heavy pigment over the entire gut (Paricelinus hopliticus, Triglops sp.) to relatively light concentrations over the dorsolateral surface only (Cottidae Type 1, Cottidae Type 2). Sometimes it ap- pears in distinctive patterns (the bars in Leptocottus ar- matus, the distinctive round melanophores in Enophrys bison) although these patterns can be obscured in poorly or improperly preserved material. Most cottid larvae develop four spines on the preoper- cle. However, some forms develop only one prominent spine (Rhamphocottus richardsoni) while others develop a large number, up to ~25, of smaller spines (Artedius spp.) on the preopercular margin. A single parietal and nuchal spine may develop (Icelinus spp.) or a cluster of smaller spines (Cottidae Type 1) or no spines at all (Lep- tocottus armatus). The same situation may prevail in the posttemporal-supracleithral region. One or more post- ocular spines may develop (Hemilepidotus spp., Triglops sp.) or they may be absent (Artedius spp.). Certain meristic characters (Table 1), particularly when used in combinations, may be useful in identifying cottid larvae. While many species have between 32 and 37 vertebrae ( = myomeres), counts at the low or high end of the range may be helpful, e.g., >40 in Triglops sp. and Nautichthys oculofasciatus and <31 in Enophrys bison. The number of pelvic fin rays ranges from to 5 with 3 being the most common count. Relatively few genera have counts of 2 (e.g., Icelinus) or 5 (e.g., Paricelinus, Scorpaenichthys) . Pectoral fin ray counts range from 12 to 26 for Oregon species and overlap in many, but counts at the low or high end of the scale help to single out some species. Also useful are dorsal and anal fin counts which for Oregon species range from 5 to 18 dorsal spines, 9 to 30 dorsal rays, and 6 to 25 anal rays. Ranges of these counts for a particular species are generally 4 or less. The difference between dorsal (D) and anal (A) soft ray counts may also be of help which is usually D > A, e.g., by 3 to 5 rays in Artedius spp., Clinocottus spp., and Oligocottus spp. and >9 in Nauthichthys oculofasciatus. In some species the D and A counts are nearly equal (Leptocottus) and in some D < A (Paricelinus hopliti- cus). Other characters that may be useful include size at hatching, size at transformation, and cirri patterns. The size at hatching, e.g., —3 mm in Artedius harringtoni vs. ~12 mm in Hemitripterus americanus (Warfel and Mer- riman 1944; Okiyama and Sando 1976) and the size at transformation, e.g., ~9 mm in Enophrys bison and ~16 mm in Chitonotus pugetensis, can serve as dis- tinguishing characters. Head cirri patterns, which may be evident in older larvae, may be important in linking larvae with known juveniles. However, they appear to develop after the pelagic phase in a number of species, e.g., Icelinus spp., and may be of limited value. ORDER OF PRESENTATION To facilitate use of this guide, the larvae described herein have been grouped according to certain primary characters (Table 2). However, a formal dichotomous key is not provided as many species remain to be de- scribed and such a key could be misleading. Rather, groups are formed on the basis of the most outstanding larval characters. The two major groupings are based on preopercular spine pattern: group A, numerous (>4); group B, 4 or less. Within group A, the larvae may be further distinguished by using a combination of charac- ters including number of preopercular spines, number of ventral midline melanophores, number of parietal spines, and presence or absence of gut diverticulae. Within group B the larvae may be placed in four sub- groups on the basis of postanal pigmentation. Within these four groups, larvae can be further separated using characters as for group A and relative body shape, pigmentation, and number of myomeres. Artedius harringtoni (St arks) (Figures 1, 2; Tables 3, 4) Literature. A 4.2 mm SL larva resembling A. harring- toni was described by Blackburn (1973) as Cottid 6. Lar- vae of this species were referred to as Artedius sp. 1 by Richardson (see footnote 4), and Richardson and Pearcy (1977). Distinguishing Features. Larvae of A. harringtoni are probably 3-4 mm long at hatching. The largest pelagic specimens we collected, ~ 13-14 mm, are beginning to develop juvenile pigmentation. A dense concentration of malanophores is present over the gut. The nape is pigmented in all but a few of the smallest larvae although this pigment becomes embed- Table 2. — Groupings of cottid larvae described in this guide based on preopercular spines and pigment patterns. Ventral 1 midline Preopercular 1 melano- Parietal Vertebrae 2 Additional spines phores spines (myomeres) distinguishing characters A. Preopercular spines numerous ( >4) Artedius harringtoni 11-21 27-30 32-34 Artedius Type 2 13-23 14-24 Multiple (32-34) dorsal gut diverticula Clinocottus acuticeps 8-13 7-9 32-33 hindgut diverticula Cottidae Type 1 19 7 Multiple (32-35) Cottidae Type 2 10-11 15-17 1 (32-34) hindgut bulges Cottidae Type 3 16-19 4-5 Multiple (33) B. Preopercular spines 4 or fewer 1. Postanal pigment absent Triglops sp. 4 1 (47) dark gut 2. Postanal pigment on ventral midline only Chitonotus pugetensis 4 41-45 1 35-36 Cottus asper 4 14-17 34-39 Enophrys bison 4 11-14 1 29-31 deep bodied Icelinus spp. 4 10-11 1 33-39 finfold pigment Leptocottus armatus 4 9-11 35-39 pigment bars on gut Paricelinus hopliticus 4 31-32 1 42 dark gut 3. Postanal pigment primarily on dorsal and ventral midline on small larvae Hemilepidotus hemilepidotus 4 14 1 35-37 dorsal pigment interrupted Hemilepidotus spinosus 4 24-34 1 35-37 dorsal pigment continuous 4. Postanal pigment heavy over body Nautichthys oculofasciatus 4 N 3 B 40-41 large pectoral fins Radulinus asprellus 4 N 1 38-39 slender bodied Radulinus boleoides ? N ? 39-40 Rhamphocottus richardsoni 1 N 1 26-28 deep bodied Scorpaenichthys marmoratus 4 N B 35-37 Cottoid Type A N 35 globose 'Maximum number at peak of development. "From Howe & Richardson 1978 (see text footnote 3). N = not distinguishable; B = bump. Table 3. — Meristies from larvae of Artedius herringtoni. (Specimen between dashed line is undergoing notochord flexion.) [* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; N = Not examined.] Pelvic Ventral Caudal fin rays Anal Pectoral fin Preoper- midline _ Dorsal Ventral Vertebrae Branchi- Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays 3.0 29 N N N N N N N 4.7 12 27 N N N N N N N 6.9 16 30NNNNNNN 7.3 -17-** 13 14 14 16 26 N N N N N N 7 9.3 IX 16 13 14 14 1,3 21 22 N N N N N N 7 11.2* IX 17 14 14 14 1,3 21 N 5 6 6 3 11 23 7 13.6 IX 17 13 15 15 1,3 18 23 N N N N N N 7 13.6* IX 17 13 14 14 1,3 11 N 4 6 6 3 12 22 7 Table 4.- -Measurements (mm) of larvae of Artedius harringtoni. (Specimen between dashed line is undergoing noto- chord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length ' Parietal spine length 3.0 4.7 0.66 0.82 0.16 0.20 0.32 0.39 1.3 1.6 0.72 1.1 0.12 0.31 N 2 N - 6.9 1.8 0.39 0.58 3.3 1.6 0.47 N - 7.3 9.3 13.6 2.0 2.0 4.3 0.47 0.35 1.1 0.78 0.94 1.1 3.4 4.5 6.6 2.4 3.0 3.8 0.90 2.0 3.6 N N N — 'Multiple spines develop along preopercular margin instead of usual 4. 5 2 N = Not measured. Artedius harringtoni 3.0mm NL 4.7mm NL 6.9mm NL Figure 1. — Larvae of Artedius harringtoni. Arfedius harringtoni ! ) /' 7.3mm SL 9.3mm SL 13. 6mm SL Figure 2. — Larvae of Artedius harringtoni. ded in later stages, >9 mm. A row of >20 melanophores (often 20-30 but >25 in larvae <6 mm) occurs along the ventral midline beginning immediately behind the anus. The number of ventral midline melanophores helps to distinguish A. harringtoni from Artedius Type 2. The posterior half of these ventral melanophores characteris- tically extend as dashes of pigment out onto the finfold particularly in larvae >7 mm. Some pigment is added to the head region and pectoral fin base in the largest specimens >12 mm. The preopercular margin develops a series of small spines which begin to appear in larvae ~4. 5-5.0 mm. These increase in number to ~ 20-25 in postflexion lar- vae. The spines on the dorsal edge and at the posterior angle of the preopercle become the largest in the series. The number of preopercular spines becomes reduced in the largest specimens, possibly due to fusion or over- growth by tissue. Artedius harringtoni never develops pronounced multiple spines in the parietal and post- temporal region as does Artedius Type 2 although mi- nute serrations are sometimes visible and a bump develops in the parietal region in larger specimens. Larvae of A. harringtoni are stubby in shape and often have a humped appearance in the nape region. The hindgut trails below the body. Snout to anus length is generally <50 c c 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 33 c o SL. The length of the pectoral fin in- creases from 4 to 26 c o SL during larval development. Artedius harringtoni is the only species of Artedius with seven branchiostegal rays (Howe and Richardson foot- note 3). Seven branchiostegals were countable in all lar- val specimens examined (25) down to ~7 mm. The inner- most pair of branchiostegals is extremely fine and small in relation to the other six and may not be formed or indistinguishable on unstained specimens under 7 mm. The adult complement or dorsal, anal, pectoral, and pelvic fin rays or ray elements can be counted in larvae ~10 mm. Relative abundance of adults in the area of larval cap- ture was also helpful. Artedius harringtoni is one of the three most common species of Artedius in Oregon tidepools, together with A. lateralis and A. fenestralis (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. har- ringtoni. Within these three species, only A. harringtoni has preorbital cirri and only A. harringtoni and A. lateralis have nasal cirri (Bolin 1944). Artedius har- ringtoni has 1 small cirrus at the base of each nasal spine while A. lateralis has 1 or 2 cirri at the base of each spine. The largest specimens in our series have a single nasal cirrus at the base of each spine and the largest specimen has small bumps in the preorbital region which may be developing cirri. Artedius Type 2 (Figures .3, 4; Tables 5, 6) Literature. Blackburn (1973) briefly described and illustrated (8.5 mm SL) larvae resembling our Artedius Discussion. Identification of this series as A. har- ringtoni was based primarily on meristic characters, es- pecially the presence of seven branchiostegal rays. K. Howe, Ph.D. candidate, Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR 97331, pers. commun. September 1978. Table 5. — Meristics from larvae of Artedius Type 2. (Specimen between dashed line is undergoing notochord flexion.) [* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; B = Bases only; N = Not examined.] Anal Pectoral Pelvic fin Preoper- Ventral midline Caudal fin rays Dorsal Ventral Vertebrae Branchi- Body length Dorsal fin Spines Rays fin rays fin rays Left Right spine & rays cular spines melano- phores Second- ary Prin- cipal Prin- cipal Second- ary Abdom- inal Cau- dal ostegal rays 3.0 4.7 — — — — N — N — ~5 24 14 N X N N N X N X N N N N N X 6.0 - - - N - 13 -15 N N N N N N N 7.3 -14B-** 12B — N — 21 15 N N N N X X 6 9.9 VIII 18 12 15 N 1,3 23 18 N X N N X N 6 11.5* IX 16 12 15 15 1,3 24 X — 6 4 — 12 22 6 11.8 IX 17 13 15 N 1,3 13 13 X N N N N N 6 Table (i. -Measurements (mm) of larvae of Artedius Type 2. (Specimen between dashed line is undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length" 3.0 4.7 0.66 1.2 0.18 0.12 0.30 0.44 1.2 1.9 0.58 1.2 0.22 0.20 N 3 X N X 6.0 1.7 0.35 0.56 2.8 1.5 0.31 N N 7.2 9.9 11.8 2.0 3.0 4.1 0.31 0.48 0.64 0.66 0.94 1.13 3.5 4.6 5.6 2.1 2.8 3.0 0.86 1.7 3.4 N N N N N N "Multiple spines develop along preopercular margin instead of usual 4. "Multiple spines develop in region of parietal spine. 3 N = Not measured. Artedius Type 2 3.0mm NL 4.7mm NL 6.0mm NL Figure 3.-Larvae of Artedius Type 2. The second figure from the top is a dorsal view showing gut diverticula. 9 Artedius Type 2 7.2mm SL 1.8mm SL Figure 4. — Larvae of Artedius Type 2. Type 2 which he called Cottid 4, "Water Wings." Eldridge (1970) also described and illustrated (3.2 mm) similar appearing larvae with gut protrusions called Cot- tid No. 4. White (1977) illustrated (3.9 mm NL) and briefly described similar larvae as Cottid III with two "horn-like extensions protruding . . . from the gut." Young stages of two known species, A. lateralis and A. fenestralis, have been described which somewhat resem- ble our Artedius Type 2, although their similarity cannot be confirmed on the basis of the literature alone (see Discussion below). Budd (1940) illustrated and described eggs and larvae (4.1 mm SL) of A. lateralis. Marliave (1975) illustrated and briefly described young stages (4, 8, 11, 14 mm TL (total length)) of A. lateralis. He also discussed eggs of A. lateralis and eggs and larvae of A. fenestralis. Larvae of this form were called Artedius sp. 2 by Richardson (footnote 4) and Richardson and Pearcy (1977). Distinguishing Features. Larvae of Artedius Type 2 probably hatch at around 3-4 mm. The largest specimens we have captured pelagically are ~14 mm, taken in a neuston net 2 km off Newport, Oreg. They are beginning to develop juvenile pigmentation. 10 In larvae, melanophores are concentrated over the dorsolateral surface of the gut. Nape pigment is usually present in larvae except at the smallest sizes but becomes embedded and obscured in later stages >8 mm. A row of usually <20 (often 15-20 but <25 in larvae <5 mm) melanophores occurs along the ventral body midline posterior to the anus. The number of ventral midline melanophores helps distinguish Artedius Type 2 from A. harringtoni which usually has >20. Approxi- mately the posterior half of these melanophores extend as characteristic dashes of pigment onto the ventral fin- fold especially in larvae >7 mm. In the largest specimens, ~12-14 mm, pigment is added to the head region, at the base of the pectoral fin, to the anterior por- tion of the spinous dorsal fin, as a bar extending from the dorsal surface of the abdominal cavity near the dorsal origin of the pectoral fin to the middle of the spinous dor- sal fin, and as groups of melanophores along the lateral midline from the gut region to the caudal fin. As in A. harringtoni a series of small spines develops along the margin of the preopercle, beginning to appear ~4.5-5.0 mm. They increase in number to ~ 20-25 in postflexion larvae, — 7-11 mm, then decrease in the larg- est specimens. Those at the dorsal edge and at the pos- terior angle of the preopercle become large relative to the rest of the spines. Clusters of spines develop in the parietal region beginning with 1 or 2 spines in 6-7 mm larvae, 3 spines in 7.5-8.5 mm larvae, and more on larger specimens. By ~9-9.5 mm clusters of spines occur in both the parietal and posttemporal regions, and these re- main visible on the largest specimens examined. Such spine clusters are never obvious on A. harringtoni. Larvae of Artedius Type 2 are similar in shape to A. harringtoni being stubby and often with a hump in the nape region. The hindgut appears to trail below the body. Snout to anus length is <50% SL. A prominent feature is the presence of gut diverticula, one on each side, protruding laterally from the dorsal surface of the abdominal cavity immediately posterior to the pectoral fin. These gut diverticula, never observed in A. har- ringtoni, are most pronounced in small larvae, becoming less conspicuous as the pectoral fin develops. They are present on the largest pelagic specimens we examined. This larval character is probably lost during transforma- tion to the juvenile stage. Body depth at the pectoral fin base ranges from about 19 to 29% SL. The length of the pectoral fin increases from 4 to 29% SL during larval development. Branchiostegal rays, which could be counted on specimens >7 mm, consistently numbered 6 for all larvae in this series compared to 7 for A. har- ringtoni. The adult complement of dorsal, anal, pectoral, and pelvic fin rays or ray elements is countable by ~10 mm. Discussion. Meristics on the largest specimens indicate they may be either A. fenestralis or A. lateralis, two of the three most abundant species of Artedius off Oregon, as previously noted. Dorsal soft ray counts of 16-18 (16=6; 17=8, 18 = 2) on larval specimens in our collec- tions seem more indicative of A. fenestralis than A. lateralis which has 15-17 (Howe and Richardson footnote 3). Head cirri patterns on the largest specimens indicate they could be A. fenestralis. The largest specimens of Artedius Type 2 have postocular cirri but show no sign of forming nasal cirri. Bolin (1944) reported that A. fenestralis has no nasal cirri while A. lateralis has 1 or 2 cirri at the base of each nasal spine. This would indicate Artedius Type 2 is A. fenestralis unless nasal cirri appear later in development. The largest specimens of A. har- ringtoni, similar in size to the largest specimens of Artedius Type 2, already have nasal cirri, indicating that the absence of nasal cirri in Type 2 is real. In a brief description Marliave (1975) stated that ". . . yolk sac larvae of Artedius fenestralis are identical to the same stage of A. lateralis except that they are perhaps a bit smaller. By the onset of fin ray formation A. fenestralis larvae have a patch of melanophores on the nape but none on the head, in contrast to A. lateralis . . . [which] develops a patch of melanophores in the oc- cipital region (none on the nape)." He further stated that A. lateralis larvae "develop lateral extensions . . . over the gut" that "distend the body wall over the dorsal in- sertion of the pectorals." Presumably then both A. lateralis and A. fenestralis larvae have gut diverticula. [Budd (1940) did not show gut diverticula on larvae of A. lateralis, but his specimens may have been too small to have developed them.] However, if the presence of nape pigment is a specific character for A. fenestralis as Marliave reported, then most, if not all, of our Artedius Type 2 may be A. fenestralis. That is if the other larval characters of <20 ventral midline melanophores, pres- ence of gut diverticula, and a pronounced cluster of spines in the parietal region are adequate to distinguish Artedius Type 2 at the species level. Interestingly, larvae of this type occur off southern California (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 confirma- tion of our Type 2 awaits identification of larvae of all Artedius species reported to occur off Oregon. If this series is A. fenestralis and we have also taken A. harringtoni, we would expect to have larvae of A. lateralis in our collections since adults of all three are relatively abundant in Oregon tidepools, as we have noted. Larvae of A. harringtoni and Artedius Type 2 were among the most abundant cottid larvae in our collec- tions. Explanations for missing A. lateralis larvae could be several: 1) they are mixed in with one of our series, e.g., Artedius Type 2; 2) they are one of our unidentified larval types and not as abundant in our collections; 3) they exhibit a different early life history strategy, e.g., they are more benthic or coastal in nature and thus would not be found in plankton collections taken 2 km or more offshore. E. H. Ahlstrom, Senior Scientist, Southwest Fisheries Center, National Marine Fisheries Service, NOAA, P.O. Box 271, La Jolla, CA 92038, pers. commun. October 1978. 11 Clinocottus acuticeps (Gilbert) (Figures 5, 6; Tables 7, 8) Literature. An 8.6 mm SL larva resembling Clinocottus acuticeps was illustrated and described by Blackburn (1973). He called his specimen Cottid 1 "Biramous anus," possibly Clinocottus sp. Eggs and larvae of two Figure 5. — Larvae of Clinocottus acuticeps. The second figure from the top is a ventral view showing hindgut diverticula. Clinocottus acuticeps 40tti 3.7 mm NL 3.7mm NL 3.9mm NL 6.9mm NL 12 Clinocottus acuticeps 7.6mm SL 16.5mm SL Figure 6. — Young of Clinocottus acuticeps. 13 other species of Clinocottus from the northeast Pacific have been described, C. analis ([Eigenmann 1892 (as Oligocottus analis); Budd 1940] including illustrations of larvae and juveniles 4.6, 5.0, 7.6, 8.3, 9.9, 10.8, 18.0, and 24.3 mm TL. Larvae of C. acuticeps were called Cottidae sp. 12 by Richardson (footnote 4) and Richardson and Pearcy (1977). Distinguishing Features. Larvae of C. acuticeps prob- ably hatch at ~3-4 mm and begin to transform to juveniles ~13-14 mm. A 16.5 mm specimen collected in a tidepool is fully transformed juvenile. Pigment is moderately scattered over the dorsolateral surface of the gut. Nape pigment is present on all small larvae but becomes embedded in larger specimens. Pig- ment is also present on the head between the eyes in all but the smallest specimens which may be faded. Melano- phores are added to the head region during development. A series of usually <15 ventral midline melanophores is present on the tail beginning at about a vertical through the anal opening in the trailing hindgut. No melano- phores are on the ventral body margin anterior to that point. Melanophores in this ventral midline series are rather inconspicuous compared to other species. The pos- terior melanophores in this ventral row do not extend as dashes onto the finfold as in Artedius spp. but may ac- tually occur on the ventral finfold rather than on the body margin in larger larvae. A single melanophore per- sists near the notochord tip, eventually appearing at the middle of the caudal fin base. A few streaks of pigment may be visible on the ventral finfold near the tail tip and later on the caudal fin. On larvae nearing transforma- tion, pigment is added to the head, pectoral fin base, and onto the anterior end of the spinous dorsal fin. Recently transformed juveniles are heavily pigmented with a noticeably darkened area at the anterior end of the first dorsal fin as in adults. A series of ~10-15 small spines develops on the preopercular margin. These begin to appear at ~6-7 mm. With development, the upper spines become the most prominent. The number of preopercular spines de- creases during transformation with only 1 dorsal spine visible in a 19.4 mm stained juvenile. No additional head spines are obvious in larvae. Parietal and nuchal spines are never apparent although the flabby skin could pos- sibly obscure their presence on unstained larvae. Larvae of C. acuticeps have an unusual gut, the poste- rior portion of which trails well below the body. Near the anus two diverticula or protrusions are evident through- out the larval period, but they are no longer visible on transformed juveniles. Snout to anus length is relatively long for cottid larvae, ranging from ~62 to 71% SL during the larval period. The larvae have a rather flabby appearance at times seeming to be encased in a bubble of loose outer skin, particularly in the head region. Body depth ranges from ~22 to 33% SL during larval develop- Table 7. — Meristics from young of Clinocottus acuticeps. (Specimen between dashed line is undergoing notochord flexion.) (* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; B = Bases only; D = Damaged; N = Not examined.] Pelvic Ventral Caudal fin ravs Dorsal fin Anal fin Pectoral fin rays fin spine Preoper- cular midline melano- Dorsal Ventral Vertebrae Abdom- Cau- Branchi- Body Second- Prin- Prin- Second- ostegal length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays 3.7 — — — — N — — 7 N N N N N N N 3.9 — — — — N — — 9 N N N N N N N 6.9 -22B-** 11B 14 N - 11 8 N N N N N N N * 7.6 VII 15 11 14 N Buds 11 7 N N N N N N N 10.2* VIII 15 12 14 14 1,3 11 N 4 6 5 2 10 14 6 10.4 VIII 16 12 1 1 N 1,3 13 7 N N N N N N N 13.8 IX 15 11 14 N 1,3 11 4 N N N N N N N 14.2* IX 16 13 14 14 1,3 12 N 7 6 5 5 10 22 6 19.4* D D 12 14 14 1,3 1 N D D D 1) 10 23 6 Table 8. — Measurements (mm) of larvae of Clinocottus acuticeps. (Specimen between dashed line is undergoing noto- chord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length 1 Parietal spine length 3.7 3.9 0.90 0.88 0.14 0.16 0.38 0.38 2.3 2.4 0.90 0.86 0.26 0.40 N" N - 6.9 2.0 0.39 0.62 4.6 2.1 0.82 N - 7.6 10.4 13.8 2.3 2.9 1 2 0.47 0.58 0.86 0.70 0.94 1 2 5.4 7.0 9.0 2.5 2.6 4.2 1.2 1.6 4.1 N N N - Multiple spines develop along preopercular margin instead of usual 4. 14 "N = Not measured. ment. The adult complement of fin rays (or elements) can be counted in larvae by ~7-8 mm. Late stage larvae have relatively long pectoral fins, ~30% SL. On larger larvae, ~13-14 mm, the inner pelvic fin ray is noticeably attached to the belly by a membrane. This diagnostic character helped link the larval series to identified juveniles. Discussion. Larvae of C. acuticeps are similar to described larvae of C. analis and C. recaluus in size at hatching, ~3-4 mm, and presence of nape and head pig- ment and pigment over the gut on small larvae. (The lack of head pigment on the smallest larvae of C. acuticeps may have been caused by fading.) All three species have a series of ventral midline melanophores but those on C. analis and C. recalvus appear to be higher in number than C. acuticeps. The posterior melanophores in this series do not appear to extend out onto the finfold in C. acuticeps as reported for the other two species. Based on the illustrations, small larvae of C. analis and C. recalvus have a much shorter relative snout to anus length than C. acuticeps. Large larvae of C. recalvus develop multiple spines on the preopercle as do C. acuticeps, but they apparently do not develop the hindgut diverticula or the flabby appearance of C. acuticeps. These marked differences in body form, main- ly the gut length and hindgut protrusions of C. acuticeps could lead to questions about the relationships within the genus Clinocottus, particularly as larvae of more species become known. Cottidae Type 1 (Figure 7; Tables 9, 10) Literature. Larvae of this type have not been previously described. These larvae were called Cottidae sp. 1C by Richardson (footnote 4) and Richardson and Pearcy (1977). Distinguishing Features. The smallest larvae in our col- lections are ~4 mm. They are relatively undeveloped and probably recently hatched. Our largest specimens are ~9 mm. They have visible fin ray elements but fins are not completely developed. Transformation occurs at some larger size. The larvae are relatively un pigmented. Melanophores occur at the nape, in a rather light scattering over the dorsolateral gut surface and in a row of <10 along the ventral midline posterior to the anus. These ventral midline melanophores are well spaced beginning ~4-5 myomeres behind the anus and extending to the poste- riormost myomere. They decrease in number anteriorly with development. The preopercle develops a high number, ~19, of small spines on its margin. A cluster of tiny spines develops in the parietal region, a few of which are visible on 4 mm specimens. Another cluster of minute spines develops in the posttemporal-supracleithral region on larger specimens. The larvae are rather stubby in shape. The hindgut trails below the body. Snout to anus length is ~ 40-50% SL. Body depth at pectoral base is -23-26% SL. The snout is relatively round in appearance. Discussion. Specific or generic identification of Cot- tidae Type 1 is not possible without additional material. These larvae are related to the Artedius-Clinocottus-Oli- gottus group of cottids based on counts of the largest specimen and the presence of multiple spines on the preopercle. Based on larval characters, these larvae ap- pear to most closely resemble A rtedius spp. The clusters of spines in the parietal and posttemporal-supra- cleithral region in these larvae have been observed only in Artedius Type 2 and Cottidae Type 3. They do not oc- cur in C. acuticeps, C. recalvus (Morris 1951), or 0. maculosus (Stein 1972, 1973) the latter of which reportedly develops only 2 spines in the parietal region. Spines on the preopercle of the largest specimen of Type 1 appear to be enlarging somewhat dorsally and at the posterior angle as in A. harringtoni and Artedius Type 2. However, their development would have to be traced in Table 9. — Meristics of larvae of Cottidae Type 1. (Specimen be- tween dashed lines is undergoing notochord flexion.) Body length Dorsal fin spines Dorsal fin rays Anal fin rays Pectoral fin rays Left Right Pelvic fin spine and rays Preoper- cular spines Ventral midline melano- phores 4.2 - - - - N 1 - - 7 6.7 - — — — N - 19 7 9.0 IX 18 12 15 N Buds 19 3 'N = Not examined. Table 10.— Measurements (mm) of larvae of Cottidae Type 1 . (Specimen between dashed lines is undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length 1 Parietal spine length - ' 4.2 0.84 0.20 0.42 1.8 0.98 0.20 N 3 N 6.7 1.4 0.20 0.52 2.6 1.6 0.55 N N 9.0 2.1 0.25 0.94 4.2 2.5 1.2 N N Multiple spines develop along preopercular margin instead of usual 4. Multiple spines develop in region of parietal spine. N = Not measured. 15 Cottidae Type 4.2 mm NL 6.7mm NL ■-■\ S 9.0mm SL Figure 7.— Larvae of Cottidae Type 1. larger specimens to determine positively if the pattern is the same as in our two Artedius types. This preopercular spine pattern differs from that in C. acuticeps and C. recalvus (Morris 1951) in which the dorsalmost spine becomes the largest. The number of preopercular spines in Type 1 is relatively high, ~ 19 as in A. harringtoni and Artedius Type 2 (>20), compared with the lower numbers for C. acuticeps (10-15), C. recalvus (4-9), and 0. maculosus (7-9). The gut shape and length resemble our Artedius types. Based on the meristic compilation by Howe and Richardson (footnote 3) (Table 1), counts on our largest Cottidae Type 1 specimen, particularly the 18 dorsal soft rays, agree with only four species: A. fenestralis, A. har- ringtoni, 0. maculosus, and 0. snyderi. The larvae are not A. harringtoni based on our description and they are not 0. maculosus based on the description by Stein (1972, 1973). Identification as 0. snyderi is not possible based on Stein's (1972) description although it seems un- likely that our larvae are this species. Larvae of 0. maculosus have a very different preopercular spine pat- tern from our Type 1, and one would expect a pattern similar to 0. maculosus on 0. snyderi. Problems in iden- tifying A. fenestralis were discussed under Artedius Type 2 and cannot be resolved at this time. L6 Cottidae Type 2 (Figure 8; Tables 11, 12) Literature. Larvae of this type have not been previously described. These larvae were called Cottidae sp. 20 by Richardson (footnote 4) and Richardson and Pearcy (1977). Distinguishing Features. The smallest larvae in the series are ~4 mm and recent);- hatched. The largest specimen is 7.4 mm in which tb-j bases of dorsal and anal Cottidae Type 2 6.4mm NL 7.4mm NL Figure 8.— Larvae of Cottidae Type 2. 17 fin rays are just forming. Size at transformation is un- known. Pretlexion larvae have pigment at the nape, a rela- tively light scattering of melanophores over the dorso- lateral surface of the gut and a series of <20 but >10 ventral midline melanophores posterior to the anus. This ventral series begins near or several myomeres posterior to a vertical through the anal opening and extends to about the fourth from last myomere. Some melano- phores in this series radiate out onto the finfold. A few melanophores occur on the finfold near the tail tip. With development some melanophores are added to the head and the number of ventral midline melanophores de- creases. Melanophores were not apparent on the caudal fin base of the largest specimen, as would be expected based on the pigmentation near the tail tip of smaller lar- vae, but the specimen was badly faded. A series of approximately 10 small spines develop on the preopercular margin. The upper spine appears some- what thicker than the lower spines on the largest specimen. A tiny parietal spine is visible on the 7.4 mm specimen but no additional head region spines are ap- parent. The shape of the gut is rather distinctive with the hindgut trailing noticeably below the body, similar to Clinocottus acuticeps. Snout to anus length is around 50% SL, longer than most species. When viewed ven- trally, the wall of the hindgut bulges out on each side of the anus reminiscent of the hindgut diverticula of C. acuticeps but not as pronounced. Body depth at the pec- toral fin base is ~21-27% SL. The snout is quite rounded. The larvae have a flabby appearance, particularly around the head region, similar to C. acuticeps. Discussion. The multiple preopercular spine pattern of Cottidae Type 2 indicates this type is related to the Arte- dius-Clinocottus-Oligocottus group. Based on a com- bination of larval characters it seems to resemble Table 11. — Meristics from larvae of Cottidae Type 2. (Specimens below dashed line are undergoing notochord flexion.) Dorsal Body fin length spines Dorsal Anal Pectoral Pelvic fin fin fin fin rays spine rays rays Left Right and rays Preoper- cular spines Ventral midline melano- phores 4.0 15 10 11 17 6 Clinocottus most closely. It has a relatively low number (<15) of multiple spines on the preopercle as does C. acuticeps (10-15) and C. recalvus (5-9) (Morris 1951). The upper preopercular spine appears to be enlarging relative to the rest, as in C. acuticeps and C. recalvus. Only a tiny parietal spine is visible on the largest specimen. Two tiny spines appear in the parietal and nuchal positions in C. recalvus but only for a brief period, and none were observed in C. acuticeps, possibly ob- scured by the flabby skin. Melanophores appear on the nape and head of all C. recalvus and all but the smallest C. acuticeps and Cottidae Type 2. The lack of pigment on the head of our smallest specimens could be a result of fading. Clinocottus recalvus larvae were described from reared and freshly preserved material which often tends to have increased numbers of and more pronounced melanophores. The hindgut trails well below the body as in C. acuticeps. Bulges appear in the hindgut wall on each side of the anus similar to the more pronounced diverticula of C. acuticeps. Such protuberances were not noted for C. recalvus, perhaps on observational over- sight. Some melanophores in the ventral midline series radiate onto the finfold as in C. recalvus. This quality was not noted in C. acuticeps but possibly could be related to quality of preservation. Melanophores occur on the finfold near the tail tip, often as streaks of pigment in all three forms. These melanophores later appear on the caudal fin in C. acuticeps and C. recalvus. They were not visible on our largest Cottidae Type 2 specimen possibly because it was faded. Cottidae Type 3 (Figure 9; Tables 13, 14) Literature. Larvae of this type have not been previously described. They were listed by Richardson (footnote 4) and Richardson and Pearcy (1977) as Oligocottus sp. 1. Distinguishing Features. The smallest larvae we have taken are ~5 mm and relatively undeveloped. Our largest specimen is 12.5 mm. Most fins appear fully formed except for secondary caudal rays, but final trans- formation must occur at a somewhat larger size. These larvae have intense pigment over the head and nape even in the smallest specimens. The head pigment is more pronounced than in any of our other lightly Table 12. — Measurements (mm) of larvae of Cottidae Type 2. (Specimens below dashed line are undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 4.0 0.8 0.16 0.42 2.2 0.84 0.24 N 2 - 6.4 7.4 1.5 1.8 0.27 0.55 0.55 0.66 3.0 3.9 1.6 2.0 0.27 0.66 N N 0.10 Multiple spines develop along preopercular margin instead of usual 4. N = Not measured. 18 Cottidae Type 3 * -■* ** »'<* $ * *«\V_, • 7.5mm NL 12.5 mm SL Figure 9. — Larvae of Cottidae Type 3. pigmented forms, i.e., those with minimal body pigment. Pigment over the dorsolateral surface of the gut is also in- tense and densely concentrated, appearing as large round melanophores. The only other pigment consists of ~4-5 ventral midline melanophores near the posteriormost myomeres and a few at the base of the forming caudal fin. The preopercle develops a series of multiple spines numbering ~ 15-20. The upper spine is the largest in the 12.5 mm specimen. A cluster of spines develops in the parietal region, appearing as a double row. A similar cluster develops in the posttemporal-supracleithral region. A nasal spine is also apparent in the 12.5 mm larva. The hindgut is somewhat trailing and snout to anus length is ~48-58% SL. The hindgut bulges slightly on either side of the anus although this is not nearly as pronounced as in Clinocottus acuticeps or Cottidae Type 2. Body depth at the pectoral fin base is ~ 22-29% SL. The snout remains relatively rounded. Discussion. Additional larger specimens are needed before identification of Cottidae Type 3 can be ac- complished with certainty. Based on current knowledge 19 of larval characters, particularly the multiple preoper- cular spine pattern, it belongs with the Artedius-Clino- cottus-Oligocottus group. The number of preopercular spines is generally less than for A. harringtoni and Artedius Type 2 which have >20, and greater than for C. acuticeps, C. recalvus, and 0. maculosus which all have <15 (Morris 1951; Stein 1973). The upper spine in Cot- tidae Type 3 becomes the largest as in Clinocottus and unlike Artedius. A cluster of spines develops in the parietal region similar to Artedius Type 2 and Cottidae Table 13.— Meristics from larvae of Cottidae Type 3. (Specimens above dashed line are undergoing notochord flexion.) Ventral Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline Body fin fin fin fin rays spine cular melano- length spines rays rays Left Right and rays spines phores 6.3 - - - - - 7 5 7.5 — 19 4 12.5 IX 15 12 14 14 1,3 16 1 Table 14.- -Measurements (mm) of larvae of Cottidae Type 3. (Specimens above dashed line are undergoing noto- chord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length 1 Parietal spine length' 6.3 7.5 1.4 1.9 0.31 0.58 0.62 0.74 3.0 4.2 1.4 2.2 0.27 0.56 N 3 N N N 12.5 3.4 0.80 1.1 7.3 3.4 2.9 N N Multiple spines develop along preopercular margin instead of usual 4. 'Multiple spines develop in region of parietal spine. ' N = Not measured. Type 1 and unlike C. acuticeps, C. recalvus, and O. maculosus. However, spines in the parietal cluster ap- pear to be fewer in number (~5) and occur in two rows in contrast to Artedius Type 2 and Cottidae Type 1. A cluster of ~4-5 spines also develops in the posttemporal- supracleithral region somewhat like Artedius Type 2 and Cottidae Type 1. The hindgut bulges slightly around the anus similar to but much less pronounced than in C. acuticeps and Cottidae Type 2. The hindgut trails more than in Artedius but less than in C. acuticeps and Cot- tidae Type 2. Cottidae Type 3 has fewer ventral midline melanophores than Artedius or Clinocottus, and these appear only along the posterior portion of the tail as reported for O. snyderi (Stein 1972). Meristics of the largest Type 3 specimen fit a number of species of Artedius and Clinocottus as well as Oligo- cottus maculosus (Table 1). Our specimens differ from larvae of the latter species described by Stein (1972, 1973) in having 1) ~19 preopercular spines instead of 7-9, 2) multiple spines in the parietal region instead of 2, and 3) only 4-5 ventral midline melanophores instead of 11- 20. Thus they must be either an Artedius or a Clino- cottus species. Triglops sp. (Figure 10; Tables 15, 16) Literature. Larvae identified as Triglops sp., which resemble our larvae, were illustrated (8.3, 12 mm SL) by Blackburn (1973) but no descriptive information was given. Larvae of other species of Triglops which have been described include 10-18 mm T. pingeli (Ehren- baum 1905-1909; Koefoed 1907 [cited by Khan 1972]; Rass 1949) from the Atlantic Ocean and Barents Sea and 8.4, 11.6, 18.9, 23.4 mm TL T. murrayi (Khan 1972) from the Atlantic Ocean. Distinguishing Features. Our smallest preflexion larva is 6.9 mm and relatively undeveloped. The largest Table 15. — Meristics from larvae of Triglops sp. (Specimens above dashed line are preflexion, that below is postflexion.) Ventral Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline Body fin fin fin fin rays spine cular melano- length spines rays rays Left Right and rays spines phores 6.9 ————— — — 7.4 _ _ _ _ _ _ _ 15.4 X 31 30 16 16 1,3 4+4 Table 16. — Measurements (mm) of larvae of Triglops sp. (Specimens above dashed line are preflexion, that below is postflexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 6.9 7.4 1.9 1.8 0.20 0.20 0.90 0.86 2.6 3.1 1.5 1.4 0.39 0.39 — - 15.4 4.4 0.80 1.4 6.8 3.2 2.9 0.62 BR 1 'BR = Broken. 20 Trig lops sp. 6.9mm NL 15.4mm SL Figure 10. — Larvae of Triglops sp. specimen we have is 15.4 mm. It has a large complement of fin rays or elements although the fins do not appear to be fully formed and transformation probably takes place at a somewhat larger size. We have taken only four specimens. The larvae have a distinctive, heavily pigmented gut, and postanal ventral midline melanophores are absent. Pigment occurs over the head and nape region of the smallest larvae, but the nape pigment is no longer visible on the largest specimen. Pigment also occurs on the pec- toral fin base. The rest of the body remains un- pigmented. Head spines on the largest specimen include a double row (anterior and posterior series) of 4 preopercular spines, a parietal spine, a nuchal spine, 2 spines in the posttemporal-supracleithral region, and 2 spines in the postocular region. The larvae are relatively slender with depth at the pec- toral base ranging from 19 to 22% SL. Snout to anus length ranges from 38 to 44% SL. The larvae have a high number of myomeres, ~ 45-46, which distinguishes them from most other cottids in the region. The oblique mouth, angular jaw, and pointed snout further dis- tinguish these larvae. Discussion. No species of Triglops are reported from Oregon waters although our Triglops sp. larvae were all collected 28 km off the Oregon coast in March and April. The larvae could be a product of spawning in more north- ern waters off Washington where two species of Triglops reportedly occur, T. macellus and T. pingeli (Hart 1973). But coastal currents generally flow northward during that time of year, minimizing the likelihood that the lar- vae were transported southward. The occurrence of relatively undeveloped larvae off the mid-Oregon coast indicates a species of Triglops may occur and spawn off Oregon. Although we cannot be positive that total adult com- plements of fin ray elements are formed in the 15.4 mm specimen, the counts of dorsal and anal fin rays, 31 and 30 respectively, are too high for T. pingeli [maximum of 28 in both fins based on Howe and Richardson (footnote 3)1. The counts are within the range given for T. macellus (27-31 in both fins) as are the pectoral fin ray counts of 16 (range 15-17, based on Howe and Richardson footnote 3). However, Howe and Richardson listed a count of 51 vertebrae for one specimen of T. macellus. We are able to count only 47 myomeres on our larvae, seemingly too low for T. macellus. The range of variability of vertebrae numbers is not yet known for T. macellus. If the one count of 51 is at the high end of the scale, 4 less vertebrae may be within the range of variability and our larvae could well be T. macellus. This remains to be confirmed. Alternatively, an undescribed species of Triglops occurs off the Oregon coast which seems less plausible. Larvae of T. pingeli from the Atlantic Ocean and Barents Sea presumably have a row of postanal ventral midline melanophores based on the illustrations by Ehrenbaum (1905-1909) and Rass (1949) and the discus- sion by Khan (1972) who compared them to larvae of T. murrayi which also have postanal ventral midline melanophores. In addition, T. pingeli also develops a row 21 of melanophores along the base of the second dorsal fin by the size of 10 mm and a mediolateral row by 11 mm [Koefoed 1907 (cited by Khan 1972)]. If larvae of T. pingeli from the Pacific develop the same pigment pat- tern as those from the Atlantic and Barent Sea, they would be easily distinguishable from our Triglops sp. lar- vae. Head and trunk pigment and shape of T. pingeli resemble our Triglops sp. indicating a generic similarity. Chitonotus pugetensis (Steindachner) (Figures 11, 12; Tables 17, 18) Literature. Larvae of this species have not been previously described. Figure 11. — Larvae of Chitonotus pugetensis. Chitonotus pugetensis 3.0mm NL 6.3mm NL 8.5mm SL 22 Chitonotus pugetensis II. 5mm SL 5.4mm SL 16. 6mm SL Figure 12. — Young of Chitonotus pugetensis. Table 17. — Meristics from larvae of Chitonotus pugetensis. (Speci- mens above dashed line are preflexion, those below are postflexion.) Ventral Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline Body fin fin fin fin rays spine cular melano- length spines rays rays Left Right and rays spines phores 3.0 — 45 6.3 — 41 8.5 -15-' 14 — — 4 37 11.5 IX 16 16 16 16 4 24 15.4 X 16 16 18 18 1,3 4 26 16.6 X 15 15 17 17 1,3 4 24 'Dorsal fin spines and soft rays cannot be distinguished. Distinguishing Features. The smallest larvae we have taken are ~3 mm and probably recently hatched. The largest pelagic specimen, 16.6 mm, is beginning to trans- form to the juvenile stage as evidenced by increasing pigmentation. The dorsolateral surface of the gut is only moderately pigmented. Additional pigment distinctively lines the ventral margin of the abdominal cavity. Pigment occurs on the head by 8.5 mm and is added to the head region through later development. A series of melanophores lines the ventral body margin postanally. These ventral midline melanophores always number >20 and usually >40 in preflexion larvae <6 mm. This is the highest number of ventral midline melanophores for preflexion 23 Table IS — Measurements (mn of larvae of Chitonotus pugetensis. (Specimens above dashed line are below are postflexion.) preflexion, those Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 3.0 6.3 0.66 1.7 0.23 0.43 0.70 0.62 0.54 2.6 1.1 1.5 0.20 0.31 - — 8.5 11.5 15.4 16.6 2.2 3.5 4.6 6.2 0.72 0.88 1.1 1.8 0.80 1.1 1.6 1.7 4.0 5.3 7.4 7.6 2.0 2.8 3.8 3.9 0.64 16 2.6 4.0 0.12 0.23 0.31 0.40 0.04 BR' 0.12 'BR = Broken. larvae of any species described in this guide. The poste- rior melanophores in this series appear to be closer together than the anterior ones. The number and spacing of the ventral midline melanophores help distinguish small larvae. Melanophores occur along the base of the caudal fin after it develops. As transformation to the juvenile stage approaches pigment is added to the ante- rior portion of the spinous dorsal fin, in the nape region, in blotches along the lateral midline, and on the pectoral fin. The preopercle becomes armed with 4 prominent spines, some of which are visible on larvae as small as ~6 mm. With development, the upper 2 spines become larger than the 2 lower ones. Small parietal and nuchal spines develop during the postflexion period and fuse together as 1 spine. Several small spines also develop in the posttemporal and supracleithral region. These latter two sets of spines are inconspicuous on the 16.6 mm specimen. A nasal spine is visible by 15 mm. Small larvae of C. pugetensis somewhat resemble liparid larvae in shape except that they have fewer myomeres. With development, the snout becomes rather pointed in appearance and the body shape resembles that of Ice linus spp. where the body outline narrows dis- tinctively near the caudal peduncle. Body depth at the pectoral fin base remains ~ 23-25% SL. Snout to anus length ranges from ~41 to 48% SL and pectoral fin length increases to ~24% SL during the larval period. The adult complement of fin rays (except dorsal spines) can be counted in 11.5 mm larvae. The deep emargination be- tween the third and fourth dorsal fin spines, a diagnostic character for C. pugetensis, is noticeable by 11.5 mm. Discussion. Chitonotus is a monotypic genus in the northeast Pacific. The larvae are rather distinctive, and with the aid of the above characters, they are not easily confused with other forms off Oregon. Cottus asper Richardson (Figure 13; Tables 19, 20) Literature. Larvae of Cottus asper were briefly de- scribed and illustrated (5.5, 9.0, 10.8 mm TL) by Stein (1972). We did not review literature on larvae of strictly freshwater species of Cottus. Distinguishing Features. Larvae hatch ~5 mm and begin to develop juvenile pigmentation ~10 mm as evi- denced by an increase in the number of melanophores in the head region. Cottus asper larvae are relatively lightly pigmented. A few melanophores cover the dorsal surface of the gut and several elongate melanophores line the ventral margin of the abdominal cavity and throat. About 15-20 evenly spaced melanophores occur along the ventral midline of the tail beginning near the anus. These decrease in number during development while a few melanophores are added laterally over the gut and one appears at the ventral edge of the preopercle. As transformation begins more pigment is added to the head, base of pectoral fin, over the gut, and along the base of the caudal fin. Table 19. — Meristics from larvae of Cottus asper. (Specimen above dashed line is preflexion, those below are postflexion.) Body length Dorsal Dorsal fin fin spines rays Anal fin rays Pectoral Pelvic fin fin rays spine Left Right and rays Preoper- cular spines Ventral midline melano- phores 5.2 - - - - N 1 - - 17 8.2 9.9 -19-' IX 18 16B 3 16 19 N 18 4 4 14 8 N = Not examined. "Dorsal fin spines and soft rays cannot be distinguished. ' B = Bases only. Table 20. — Measurements (mm) of larvae of Cottus asper. (Specimen above dashed line is preflexion, those below are postflexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 5.2 0.9 0.12 0.36 2.1 0.78 0.51 - - 8.2 9.9 1.9 2.6 0.35 0.66 0.55 0.74 3.8 4.2 1.4 1.8 0.62 2.5 -0.03 0.33 - 24 Cottus osper 5.2mm NL 8.2mm SL 9.9mm SL Figure 13. — Larvae of Cottus asper. Four pronounced spines develop on the preopercular margin, evident in 8 mm larvae. No additional head spines form with development. The shape of C. asper larvae is rather distinctive, being somewhat blennioid in appearance. This is partly related to the characteristic coiling and shape of the gut which is unlike any of the other cottids described here. Snout to anus length is usually between 40 and 46% SL. Body depth at the pectoral fin base is slender relative to body length, usually <20% SL. Total fin rays or elements are countable by ~10 mm. Discussion. Two species of Cottus, C. asper and C. aleuticus, are found in brackish water off Oregon. Only larvae of C. asper have been identified in coastal plank- ton collections. Ringstad 8 provided evidence that in a British Columbia coastal stream most C. asper spawn in the estuary while C. aleuticus spawn primarily in fresh water. Thus larvae of C. aleuticus may simply not occur "Ringstad, N. R. 1974. Food competition between freshwater sculpins (Genus Cottus) and juvenile coho salmon {Oncorhynchus ki- sutch): an experimental and ecological study in a British Columbia coastal stream. Environ. Can., Fish. Mar. Serv., Tech. Rep. 457, 88 p. in coastal plankton. Larvae of C. aleuticus have not been described. Meristics, especially anal (usually >15 for asper and <15 for aleuticus) fin ray numbers will gen- erally separate the two (Scott and Crossman 1973; Howe and Richardson footnote 3). Enophrys bison (Girard) (Figure 14; Tables 21, 22) Literature. Blackburn (1973) illustrated and briefly de- Table 21. — Meristics from larvae of Enophrys bison. (Specimen be- tween dashed lines is undergoing notochord flexion.) Body length Dorsal fin spines Dorsal fin rays Anal Pectoral fin fin rays rays Left Right Pelvic fin spine and rays Preoper- cular spines Ventral midline melano- phores 4.8 - - - - N' - - 14 7.0 - - — — N — 1 . 11 9.1 IX 11 9 16 N [,'i 4 10 N = Not examined. 2. r > Enophys bison 4.8mm NL 7.0mm NL 9.1mm SL Figure 14. — Young of Enophrys bison. 26 Table 22. — Measurements (mm) of larvae of Enophrys bison. (Specimen between dashed line is undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 4.8 1.1 0.2 0.55 2.3 1.1 0.37 - - 7.0 2.0 0.5 0.66 3.8 1.9 0.55 0.35 0.27 9.1 3.4 0.6 0.90 5.0 3.4 2.08 0.43 0.27 scribed a 7.5 mm larva of E. bison and Marliave (1975) a 10 mm TL larva. Misitano (1978) described a develop- mental series, including eggs, of E. bison (5.0, 5.4, 5.8, 6.7, 7.1, 7.6 mm SL) based on laboratory-reared material. Distinguishing Features. Larvae are ~5 mm at hatching. The addition of juvenile pigment may begin in specimens as small as 7 mm in the laboratory but may occur later in the natural environment based on our material. Transition from pelagic to benthic habitat begins in specimens as small as 7.6-7.8 mm in the labo- ratory but Marliave (1975) captured a 10 mm TL trans- forming specimen in the plankton and we took trans- forming specimens up to ~9.5 mm in the neuston. Even with this variation, transformation to juveniles takes place at a relatively small size in E. bison compared to other known cottids. Pigmentation over the dorsolateral surface of the gut is characteristic and intense. Melanophores often appear as tightly packed discrete circles. Additional melanophores are on the ventrolateral surface of the gut of fresh mate- rial but are faded on our smallest specimens. Melano- phores are present on the nape, appearing very dark in freshly preserved specimens. The number of postanal ventral midline melanophores is usually <15. A dis- tinctive space of several myomeres in length exists between the anus and the anteriormost ventral melano- phore. The ventral midline melanophore series is rela- tively inconspicuous, compared to other cottid species. During development, pigment is added to the head which becomes entirely covered with melanophores by transformation. Melanophores are added beneath the spinous dorsal fin forming a band extending to the gut, to the base of the pectoral fin, to the ventrolateral region of the gut, and eventually beneath the second dorsal fin ex- tending as a band to the ventral body margin. A few melanophores appear at the base of the caudal fin. Larvae of E. bison develop very prominent head spines. Four pronounced preopercular spines are visible on all but the smallest larvae. The length of the second preopercular spine is 18% HL in a specimen undergoing notochord flexion. They also develop a rather large parietal spine, visible by ~6 mm. With development, a nuchal spine appears posterior to the parietal and fuses with it, and another spine develops on the parietal ridge anterior to the parietal spine. A postocular spine develops over the eye. A spinelike bump develops in late larvae in the posttemporal-supracleithral region and a nasal spine appears on the snout. Two spiny tips (mid- opercular and interopercular-subopercular) can be seen on the gill cover margin. All but the smallest larvae are relatively deep bodied with body depth at the pectoral fin base increasing to ~37% SL on our transforming specimens. Snout to anus length increases from ~48 to 55% SL with development. The preanal finfold described by Misitano (1978) in lar- vae <6.6 mm was not visible on our specimens probably because of poor condition. All fin rays are countable by the onset of the transformation period at ~9 mm in our material. Pectoral fin length reaches 23% SL by trans- formation. Enophrys bison larvae have a character- istically low number of myomeres (vertebrae = 29-31) (Howe and Richardson footnote 3) which helps to distin- guish them from other cottids off Oregon most of which have >31 vertebrae (Table 1). Discussion. Only one species of Enophrys occurs off Oregon and the larvae are rather distinctive and not like- ly to be confused with other species. However, two or three additional species of Enophrys may occur in the northeast Pacific, E. diceraus, E. lucasi, and E. taurina (Sandercock and Wilimovsky 1968; Quast and Hall 1972). Their larvae have not yet been described. Whether larval characters useful for E. bison (i.e., distinctive gut pigment, intense nape pigment, pronounced head spines, gap between anus and first ventral midline melano- phore, small size at transformation) would be helpful in distinguishing larvae of the other species remains to be determined. Certainly the low number of vertebrae (26- 29) of E. taurina (Howe and Richardson footnote 3) will be a useful character. Larvae of E. bubalis and E. lillje- borgi (Russell 1976) closely resemble E. bison with some differences in pigmentation and spination. Icelinus spp. (Figures 15, 16, 17; Tables 23, 24) Literature. A 4.3 mm larva resembling this type was illustrated and described by Blackburn (1973) as Cottid 3. These larvae were referred to as Icelinus sp. 1 by Rich- ardson (footnote 4) and Richardson and Pearcy (1977). Distinguishing Features. The smallest larvae in our col- lections are ~3-4 mm and probably recently hatched. The largest specimens we have collected pelagically are 27 Icelinus spp. 3.3mm NL 8.6mm NL V— ^o.. * >: 10.9mm SL Figure 15. — Larvae of Icelinus spp. ~ 17-18 mm, some taken in neuston tows, and they are beginning to develop juvenile pigmentation. Larvae up to ~8-9 mm are characterized by a series of large melanistic spots on the dorsal and ventral finfold, when intact. Most all of our specimens have 2 spots dor- sally and ventrally but one had 3 ventral spots, and Blackburn's (1973) specimen had 3 spots dorsally and ventrally. Whether the number of finfold spots is a specific character or merely individual variation is not known at this time. Moderate pigment occurs over the dorsolateral surface of the gut and some melanophores are present near the base of the cleithrum. A series of dis- tinctive melanophores, usually <15, occurs along the ventral midline of the tail beginning immediately behind the anus. These melanophores vary in size with some of them appearing more pronounced than others. One or several melanophores clustered at the posterior end of this ventral row distinctively appear at the ventral margin of the caudal fin base after it is formed. With development pigment is added to the head and over the gut with little additional change. Some larger speci- mens, >12 mm, have 2 distinct melanophores on the dor- sal midline in the tail region. Whether this is a specific character is unknown, but we have been unable to link a complete developmental series together based on the presence of these dorsal melanophores. Some speci- mens, with and without dorsal midline melanophores, have an additional distinct pigment spot on the dorsal- most rays of the caudal fin near its base. In specimens nearing transformation, pigment increases over the head region, on the spinous dorsal fin, and on the pectoral fin base. 28 Icelinus spp. 13. 8mm SL 15. 2mm SL 6.5mm SL Figure 16. — Young of Icelinus spp. Spines are first noticeable on the preopercle ~8-9 mm and 4 are obvious by 10-11 mm. The upper one becomes somewhat larger than the rest. Parietal and nuchal spines develop and fuse together. Several spines appear in the posttemporal-supracleithral region eventually be- ing reduced to bumps. These are difficult to distinguish from developing spinelike scales along the lateral line and below the spinous dorsal fin. A nasal spine is obvious on the largest specimens. The gut shape is rather distinctive in larvae of Ice- linus spp. Snout to anus length is relatively short in small larvae, <40% SL, but increases to ~50% SL on larger specimens. Body depth at the pectoral fin base is -20% SL in small larvae and -25-30% SL on older lar- vae. Larvae of Icelinus spp. have a very characteristic body shape. Body depth appears to be constricted just behind the anus, bulges out slightly in the midtail region, and then narrows distinctively again near the tail tip or caudal peduncle. They also develop a rather pointed snout which is apparent on larvae >8 mm. Pectoral fin length is ~ 25-30% SL on the largest specimens. The adult complement of fin rays or elements is countable on specimens >12 mm including the diagnostic pelvic fin count of 1,2. Discussion. Identification of the largest specimens to the genus level was relatively easy since Icelinus is the only representative off Oregon except Zesticelus pro- fundorum with 2 pelvic fin rays. The distinctive body shape and pigmentation helped link the larval series to- 29 Icelinus spp. ^ ^^^^ 12. 5mm SL 16.6mm SL Figure 17. — Young of Icelinus spp. Table 23. — Meristics from larvae of Icelinus spp. (Specimen between dashed line is undergoing notochord flexion.) [* = Stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; t = Form with two melanophores on dorsal midline; N = Not examined.] Pelvic Ventral Caudal fin rays Anal Pectoral fin Preoper- midline _ Dorsal Ventral Vertebrae Branchi- Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal length Spines Rays rays Left Right & rays spines phores ary cipal cipal ary inal dal rays 3.3 10NNNNNNN 5.1 11 N N N N N N N 8.6 3 10 N N N N N N N 10.9 -25-** 12 15 15 4 10 N N N N N N N tl2.5 X 16 12 16 16 1,2 4 8 N N N N N N N 13.8 X 16 12 16 16 1,2 4 10 N N N N N N N 1 14.1 X 16 12 16 16 1,2 4 11 N N N N N N N 14.4* XI 15 12 15 15 1,2 4 N 5 6 6 4 11 23 6 15.2 X 16 12 15 15 1,2 4 11 N N N N N N N U5.5* X 16 12 15 15 1,2 4 N 6 6 6 4 11 23 6 16.5 X 16 12 16 15 1,2 4 10 N N N N N N N U6.6 X 16 13 15 15 1,2 4 12 N N N N N N N gether at least at the generic level. However, specific identification is complicated because most meristics of the four Icelinus species reported to occur off Oregon, /. burchami, I. filamentosus, I. oculatus, and /. tenuis overlap. An anal fin ray count of 12 agrees only with /. burchami from Oregon waters (Table 1), a species which is reportedly rare (Howe and Richardson footnote 3). Four other northeast Pacific species of Icelinus (I. borealis, I. cavifrons, I. fimbriatus, and /. quadri- seriatus) can have an anal ray count of 12 but none reportedly occur off Oregon (Howe and Richardson foot- note 3). Characters used to distinguish adults such as cir- ri patterns and elongated dorsal fin spines are not developed on our largest pelagic specimens. These fac- tors together with inability to find consistent larval char- acters to split our Icelinus spp. larvae into subgroups po- tentially equivalent to species forces us to keep our indentification at the generic level. 30 Table 24. — Measurements (mm) of larvae of Icelinus spp. (Specimen between dashed line is undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 3.3 5.1 0.76 0.86 0.10 0.12 0.32 0.39 1.2 1.7 0.68 1.0 0.26 0.31 - — 8.6 1.8 0.51 0.55 3.4 1.7 0.39 - - 10.9 n.2.5 13.8 U4.1 15.2 16.5 16.6 2.5 4.0 4.2 5.2 5.2 7.2 6.1 0.80 1.0 1.0 1.4 1.2 2.7 1.8 0.88 1.2 1.3 1.4 1.5 1.4 1.4 4.6 6.2 6.4 7.0 7.7 9.0 8.3 2.2 3.0 3.6 4.2 4.0 4.8 4.2 1.04 2.2 2.4 3.0 4.1 3.8 3.2 0.22 0.32 0.55 0.48 0.56 0.56 0.32 0.12 0.26 0.31 0.24 0.24 0.24 0.32 + Form with 2 melanophores on dorsal midline. Leptocottus armatus Girard (Figure 18; Tables 25, 26) Literature. Eggs and recently hatched larvae (~4 mm TL) were illustrated and described by Jones (1962). Eggs and young (8, 12, 13 mm TL) were briefly discussed by Marliave (1975) and larvae by Blackburn (1973). White Figure 18. — Larvae of Leptocottus armatus. Leptocottus armatus 5.1mm NL i:;::y 8.1mm NL ll.lmm SL 31 Table 25. — Meristics from larvae of Leptocottus armatus. (Specimen between dashed line is undergoing notochord flexion.) Body length Dorsal fin spines Dorsal fin rays Anal fin rays Pectoral fin ravs Left Right Pelvic fin spine and rays Preoper- cular spines Ventral midline melano- phores 5.1 - - - - N 1 - - 11 8.1 — — 15 - N - 4 9 11.1 VIII 18 17 19 N — 4 7 Discussion. Leptocottus is a monotypic genus in the northwest Pacific. The larvae with their characteristic bars of pigment over the gut should not be easily con- fused with any other species unless damage to the speci- men results in a blurring of the gut pigment. Even so, the internal pigment on the snout serves as a useful diagnostic character. N = Not examined. Table 26. — Measurements (mm) of larvae of Leptocottus armatus. (Specimen between dashed lines is undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 5.1 0.92 0.18 0.25 2.0 0.84 0.23 - - 8.1 1.9 0.31 0.32 3.2 1.6 0.78 0.10 - 11.1 2.8 0.62 0.82 4.9 2.3 2.2 0.21 — (1977) illustrated (4.9 mm NL) and briefly described small larvae. Blackburn's Cottid 5 (5.8 mm SL) was probably this species as was Eldridge's (1970) Cottid No. I (5.1 mm). Distinguishing Features. Hatching takes place at 3.9- 4.8 mm TL. Our largest pelagic specimen is 11 mm and may be nearing transformation as indicated by in- creased pigmentation over the head. Marliave's (1975) 13 mm TL specimen was transformed and had assumed a benthic habit. Larvae have distinctive pigment over the dorsolateral surface of the gut which always appears as ~6-8 bars ex- cept in damaged specimens. No other known cottid lar- vae have such gut pigment. Larvae also have distinctive internal pigment on the snout at the anterior part of the forebrain and extending posteriorly from the eye to the pectoral fin. The latter becomes obscured by muscle but the former remains visible throughout the larval period. Several melanophores occur along the ventral margin of the gut cavity. A row of usually <15 rather prominent melanophores occurs along the ventral midline of the tail beginning about 5 myomeres posterior to the anus. With development, melanophores are added to the head and a few are present at the base of the caudal fin. Four prominent spines develop along the preopercular margin, evident in ~7 mm larvae. No additional head spines are formed. Larvae are rather slender with body depth at pectoral fin base comprising 21% SL in the largest specimen. Snout to anus length increases slightly from 39 to 44% SL in the specimens examined. The snout has a rounded appearance never becoming pointed. The adult comple- ment of dorsal, anal, and pectoral fin rays is countable by II mm; however, the pelvic fins are not developed at this time. Paricelinus hopliticus Eigenmann and Eigenmann (Figures 19, 20; Tables 27, 28) Literature. Larvae have not previously been described. Distinguishing Features. The smallest larvae in our col- lections, ~5-6 mm appear to be recently hatched. An 18.6 mm specimen is beginning to transform as evi- denced by increased head pigmentation and body spi- nation (modified scales). A 25.6 mm specimen captured pelagically appears to be a fully transformed juvenile. Larvae have a distinctively pigmented gut with mela- nophores covering the entire abdominal cavity. Pigment is scattered over the head region. Over 30 melanophores line the ventral body midline in the tail region beginning just behind the anus in small larvae but the number de- creases to ~ 15-20 in postflexion stages. At the posterior end of this ventral row a separate group of distinctive melanophores occurs near the tail tip extending onto the finfold. These melanophores line the base of the caudal fin after it forms. Little pigment is added during the lar- Table 27. — Meristics from young of Paricelinus hopliticus. (Speci- mens above dashed line are preflexion, those below are postflexion.) Ventral Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline Body fin fin fin fin rays spine cular melano- length spines rays rays Left Right and rays spines phores 5.6 — — — — N 1 — — 32 6.2 — — — — N — — 31 13.8 XII 19 23 15 N 1,5 4 17 18.6 XII 19 23 15 N 1,5 4 17 25.6 XII 19 23 15 N 1,5 4 19 'N = Not examined. 32 Table 28. — Measurements (mm) of young of Paricelinus hopliticus. (Specimens above dashed line are preflexion, those below are postflexion.) Body depth 2d preoper- Body Head Snout Eye Snout to at pectoral Pectoral cular Parietal length length length diameter anus length fin base fin length spine length spine length 5.6 1.2 0.23 0.51 2.0 0.86 0.16 — — 6.2 1.7 0.31 0.70 2.2 1.0 0.23 — — 13.8 4.3 1.0 1.7 7.0 4.2 BR 1 BR BR 18.6 5.0 0.96 1.4 8.8 4.6 4.2 0.80 -0.24 25.6 8.6 2.1 2.7 11.5 5.3 7.4 0.96 0.48 'BR = Broken. val period except over the head and at the base of the pectoral fin. The pelagically captured juvenile has much increased pigmentation in blotches over the entire body and on most fins. Four spines develop on the preopercular margin in postflexion larvae. A postocular and a nasal spine also become prominent. Spines appear to develop in the parietal and nuchal region and also anteriorly along the parietal ridge. These are difficult to distinguish from the spiny scales that develop dorsally. Similarly, several spines appear to develop in the posttemporal-supra- cleithral region but are difficult to distinguish from the spiny lateral line scales. Three spine tips (midopercular, subopercular, interopercular) develop along the gill cover margin. A cleithral spine becomes evident in the larger specimens. Snout to anus length is relatively short in preflexion larvae, ~35-36% SL but increases with development. Body depth at the pectoral base is also small relative to body length, ~ 15-16% SL on preflexion larvae, in- Paricelinus hopliticus 5.6mm NL 6.2mm NL ;s> 13. 8mm SL Figure 19. — Larvae of Paricelinus hopliticus. 33 Paricelinus hopliticus 18. 6mm SL 25.6mm SL Figure 20. — Young of Paricelinus hopliticus. creasing to ~ 25-30% SL during the larval period then de- creasing in the juvenile stage. The general body shape of the larvae resembles that of Icelinus spp. The snout be- comes noticeably pointed. Discussion. Paricelinus is a monotypic genus in the northeast Pacific and its larvae are distinctive. The only other cottid larvae we have encountered with such a heavily pigmented gut are those of our Triglops sp. which lack ventral midline melanophores in the tail region. that work, brief descriptions of other hemilepidotine sculpins were also included: Hemilepidotus gilberti (as H. g. gilberti) (7.5, 11.4, 17.5 mm), H. jordani (6.4, 10.7, 13.0 mm), and Melletes papilio (10.7, 13.7 mm) with some comments on H. zapus (as H. g. zapus). Young of H. hemilepidotus, ~20 mm, were described by Peden (1964) who also described young (~20 mm) of H. jordani, H. spinosus, and H. zapus. Young of H. gilberti (7.1, 11.6, 19.2, 24.8, 32.5 mm) were described by Hattori (1964). Hemilepitodus hemilepidotus (Tilesius) (Figures 21, 22; Tables 29, 30) Literature. Gorbunova (1964) illustrated (7.25, 10.5 mm) and briefly described larvae ofH. hemilepidotus. In Distinguishing Features. Newly hatched, reared larvae are ~5-6 mm long (Dunn footnote 5) and the smallest larvae in our collections are about that size. By ~19-20 mm specimens have adult meristic complements and have acquired a juvenile appearance, although speci- mens as large as 23 mm have been captured pelagically Table 29. — Meristics from young of Hemilepidotus hemilepidotus. (Specimen between dashed lines is undergoing notochord flexion.) [* = Specimen stained with Alizarin Red S; N = Not examined.] Anal fin rays Pectoral fin rays Left Right Pelvic fin spine and rays Preoper- cular spines Ventral midline melano- phores Caudal fin rays Dorsal Ventral Vertebrae Abdom- Cau- inal dal Branchi- ostegal rays Dorsal scale rows Scales above Body Dorsal fin length Spines Rays Second- ary Prin- cipal Prin- cipal Second- ary lateral line Left Right 5.8 — — 5.9 — — — — N — N — — 14 14 N N N N N N N N N N N N N N — — N — N 9.1 — — N - 2 9 N N N N N N N - — N 10.7 11.4* 11.5 19.0 23.0* IX XI XI XI 19 17 18 20 19 15 12 15 16 17 17 13 17 10 16 N 13 N 10 10 Buds Buds Buds 1,4 1,4 II N 7 2 N N N 6 6 N N N N 6 6 N N 12 19 N N N N 12 24 19 27 N N 23 34 at the surface. Thus there may be a pelagic juvenile phase similar to that of rockfishes, Sebastes spp. The smallest transformed specimen collected in Oregon tide- pools was 21 mm SL. In our collection we have no speci- mens between ~11.5 and 19 mm. Newly hatched larvae have pigment on the head and along the dorsal midline of the body extending to all but the posterior 3d or 4th myomere. An unpigmented area occurs on the dorsal midline between about myomeres 4- 11 which fills in with pigment by ~6-7 mm. The dorso- lateral surface of the gut is moderately pigmented. A series of melanophores extends along the ventral midline beginning at about the 9th or 10th postanal myomere extending to all but the last 3 or 4 myomeres. Mela- nophores in this series usually number <15. With devel- opment, melanophores are added over the head, an- terodorsal body surface, and the gut. Some internal melanophores appear to extend dorsally from the ven- tral midline melanophores in larvae as small as 6 mm. Additional internal melanophores appear ventrolat- erally between the notochord and ventral midline. By 9 mm, internal melanophores can be seen in a row above Hemilepidotus hemilepidotus 5.8mm NL 5.9mm SL 9.1mm NL Figure 21.— Larvae of Hemilepidotus hemilepidotus. 35 Table 30. — Measurements (mm) of young of Hemilepidotus hemilepidotus. (Specimen between dashed lines is under- going notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 5.8 5.9 0.90 1.0 0.06 0.27 0.52 0.58 2.0 2.1 1.0 0.86 0.32 0.34 — - 9.1 2.0 0.27 0.9 3.4 1.7 0.62 0.22 0.10 10.7 11.5 19.0 2.7 3.7 6.7 0.47 0.96 1.28 1.2 1.3 2.2 4.6 6.4 11.0 2.6 3.3 6.0 1.1 1.7 4.0 0.39 0.51 0.9 0.27 BR 1 0.31 BR = Broken. Hemilepidotus hemilepidotus I0.7mm SL II. 5mm SL 19.0mm SL Figure 22. — Young of Hemilepidotus hemilepidotus. 36 and below the notochord along much of the body. These internal melanophores remain visible until the lateral musculature obscures them. External melanophores are not added laterally until larvae are >11 mm, and then as a relatively light scattering compared to H. spinosus. The ventral midline melanophores gradually become in- ternal but remain visible through the skin and serve as a diagnostic character to distinguish them from H. spino- sus until dorsal scale rows develop (see Discussion). By 19 mm the entire body except the caudal peduncle and ventral body margin are covered with pigment which is darker dorsally and lighter below. Four prominent spines develop on the preopercle, some of which are visible by 9 mm. A parietal spine can be seen at ~9 mm and later a nuchal spine, the two fusing to- gether late in the larval period. A postocular spine develops over the eye ~9 mm and remains obvious at least until 11 mm, probably disappearing during trans- formation. This spine develops only as a minute spinule for a brief period, ~8-9.3 mm in H. spinosus, barely visi- ble without staining. With development three spines ap- pear in the posttemporal-supracleithral region, being reduced to bumps in larger specimens. A cleithral spine is apparent by ~ 19-20 mm. A nasal spine develops on the snout. Tips of 3 spines (midopercular, interopercular, subopercular) can be seen on the margin of the gill cover by ~11 mm. Relative body depth increases with development from ~15-18 to 32% SL as does snout to anus distance from ~35 to 58%. Pectoral fin length increases to ~21% SL by 19 mm. Adult complements of fin rays, dorsal scale rows, and scales above the lateral line are countable in the 19 mm specimen and probably in smaller specimens based on H. spinosus in which adult complements are attained by ~14 mm. Gill membranes are noticeably free from the isthmus in the 19 mm specimen. Discussion. The relative position of the first postanal ventral midline melanophore and the posterior extent of the ventral series serve as excellent diagnostic charac- ters to separate larvae of H. hemilepidotus from H. spi- nosus until the dorsal scale rows are formed. In H. hemi- lepidotus the ventral midline melanophore series begins at about the 9th or 10th postanal myomere and consists of usually <15 melanophores. In H. spinosus this ven- tral row begins immediately behind the anus and con- sists of usually >15 melanophores in all but the largest specimens. In larger specimens, the number of dorsal scale rows separate the two species, 4 or 5 in H. hemi- lepidotus and 6-8 in H. spinosus. These can be counted in specimens as small as 14 mm. The placement and number of ventral midline mela- nophores may not serve to distinguish H. hemilepidotus from other species, e.g., H. gilberti, H. jordani, and H. zapus, based on Gorbunova's (1964) and Hattori's (1964) descriptions. All four species reportedly have the same ventral midline melanophore pattern. And small larvae of all four reportedly have an unpigmented break in the dorsal midline melanophore series which fills in with development. Additional characters given by Gorbunova do not appear to be adequate to distinguish larvae among these four species. One character used by Gorbunova to distinguish late larvae of//, gilberti, i.e., presence of a row of scales above the anal fin, is invalid. We have observed such scales in H. spinosus and H. hemilepido- tus, and Peden (1964) has shown them in young ofH. jor- dani and H. zapus. These become overgrown on larger fish. Gorbunova stated that larvae of H. gilberti develop 5 preopercular spines which would help distinguish them but this is probably an error in observation, based on our knowledge of other cottid larvae and illustrations by Hattori. Pigment patterns as reported by Gorbunova seem inadequate to distinguish species. Closer attention to detail in exact numbers of ventral melanophores and their placement with respect to myomere number to- gether with a more detailed examination of external lat- eral pigment and internal pigment along the notochord may help resolve the problems of larval identification. Meristic and morphometric characters as discussed by Peden will be useful in identifying larger specimens, > 18-20 mm. Relative body proportions may also be of some help but these must be better defined to evaluate their usefulness in young specimens. Larvae (10.7, 13.7 mm) of another hemilepidotine sculpin from the Bering sea, Melletes papilio, as de- scribed by Gorbunova (1964) are easily distinguished by the extension of the dorsal and ventral midline melano- phores onto and around the urostyle. See also Discussion under H. spinosus. Hemilepidotus spinosus (Ayres) (Figures 23, 24; Tables 31, 32) Literature. Follett (1952) illustrated (12, 21 mm SL) and briefly described several specimens of H. spinosus which we confirm. Peden (1964) also discussed young (~20 mm) of this species. (See also Literature section un- der H. hemilepidotus.) Distinguishing Features. The smallest larvae in our col- lections are ~5 mm and appear to be recently hatched. Specimens as large as 27 mm have been captured pelag- ically although most juvenile characteristics appear to be present by ~ 19-20 mm. The smallest larvae examined, ~5 mm, have pigment on the head and in a continuous line on the dorsal body margin from the head to the posteriormost myomere. Some pigment occurs on the dorsolateral surface of the gut. A row of >15 melanophores extends along the ven- tral body midline from immediately behind the anus to the posteriormost myomere. A single melanophore is usually apparent posterior to this line about midway to the notochord tip. This melanophore later appears on the midcaudal fin base between the upper and lower hy- pural elements. With development external melano- phores are added on the head, over the gut and laterally in H. spinosus larvae as small as 6 mm in contrast to H. hemilepidotus. By 8-9 mm, internal pigment is notice- able in a line above the notochord along most of the 37 Hemilepidotus spinosus 6.6mm NL 8.9mm NL Figure 23. — Larvae of Hemilepidotus spinosus. body. Internal pigment never becomes obvious below the notochord as in H. hemilepidotus, possibly because of thicker musculature or because the more intense external lateral melanophores obscure it. Pigment increases over the dorsal body surface first anteriorly in the nape region then posteriorly extending laterally until most of the body is pigmented, except ventrally and on the caudal peduncle, by ~ 19-20 mm. The ventral midline melano- phores become embedded but remain visible through the skin at least to ~14 mm, which serve to distinguish these larvae from H. hemilepidotus. Four preopercular spines develop, beginning ~7 mm. A parietal spine can be seen ~7 mm and later a nuchal spine develops posterior to it fusing with it in the largest specimens. A minute postocular spine develops over the eye in larvae 8.0-9.3 mm and then disappears. This spine never becomes prominent as in H. hemilepidotus and is barely visible without staining. Three spines develop in the posttemporal-supracleithral region but become reduced to bumps. A nasal spine and a cleithral spine are present by 19 mm. Three spine tips (midopercular, sub- .'58 Hemilepidotus spinosus II. Omm SL <■*%*,■&■**!* 1 1.8mm SL 19.0mm SL Figure 24. — Young of Hemilepidotus spinosus. opercular, interopercular) are visible on the gill cover margin by ~11-12 mm. Body depth increases from ~ 15-16% to -32-34% SL with development as does snout to anus distance, 38-39% to 60-61% SL. Pectoral fin length increases to ~26% SL by 19-20 mm. Adult complements of fin rays, dorsal scale rows, and scales above the lateral line are present by ~14 mm. In 19-20 mm specimens the gill membranes are noticeably attached to the isthmus. Discussion. Early life history strategies appear to differ between H. spinosus and H. hemilepidotus. Larvae of H. spinosus are much more abundant in our plankton sam- ples than H. hemilepidotus and they have been collected farther offshore, as far as 111 km from the coast. We have taken late larvae-early juvenile stages of both species in neuston collections but H. spinosus is much more com- mon. Collections in Oregon tidepools have yielded pri- marily young juveniles of H. hemilepidotus and only rarely H. spinosus. Our data indicate that H. hemilepi- dotus generally spawns closer to shore and the larvae are not as neustonic as H. spinosus. Tidepools may be impor- tant nursery areas for juvenile H. hemilepidotus whereas 39 Table 31.— Meristics from young of Hemilepidotus spinosus. (Specimen between dashed lines is undergoing notochord flexion.) [* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and soft rays cannot be distinguished; B = Bases only; N = Not examined] Ventral Caudal fin rays Scales Dorsal fin Anal fin Pectoral fin rays Pelvic fin spine Preoper- cular midline melano- Dorsal Ventral Vertebrae Abdom- Cau- Branchi- ostegal Dorsal scale ab later sve Body Second- Prin- Prin- Second- il line length Spines Rays rays Left Right and rays spines phores ary cipal cipal ary inal dal rays rows Left Right 5.0 24 N N N N N N N _ 5.7* — — — — — — — N — — — — — — — 6.2 29 N N N N N N N — N 6.6 34 N N N N N N N — 6.6 2 29 N N N N N N N — N 7.2 2 27 N N N N N N N — - N 7.6* _ — — — — — — N _ _ 8.0* 2 N 8.3 — — 14B — — — 3 24 N N N N N N N — N 8.6 -18B-** 15B — — — 3 31 N N N N N N N — — N 8.8* — — — 6 6 — 4 N — 4 4 — 12 18 6 — — 8.9 -18B-** 14B — — — 3 26 N N N N N N N — — — 9.1 -19 ** 15 — — — 4 27 N N X X N N N — — N 9.3* — — — 8 7 — 4 N — 4 5 — 12 21 6 — 10.1 -20 ** 15 1 I X — 4 N N X N X N N - - N 10.4* XI 19 15 15 15 1,1 4 4 6 6 3 12 24 6 10.6 -19 ** 14 14 N — 4 21 N N N N N N N — — N 11.0 XI 19 15 15 15 Buds 4 22 N N N X N N N — — — 11.0 X 20 15 15 15 Buds 4 25 N N N X N N N — — N 11.2 XI 18 15 15 15 Buds 4 21 X N N X N N N — — N 11.8* XI 19 16 15 15 1,4 4 N 7 6 6 6 VI 24 6 — — — 11.8 XI 18 15 15 15 1,4 4 22 N N X X N N N — — — 11.8 XI in 14 15 15 Buds 4 18 N N N X N N N — — N 12.0* XI 20 15 15 15 1,4 4 N 7 6 6 3 12 24 6 — — 12.6* XI 19 16 16 16 1,4 4 N 7 6 6 5 12 24 6 — — — 13.1* XI 19 16 15 15 1,4 4 N 8 6 6 7 L2 24 N 4 32 32 13.4 XI 19 15 15 15 Buds 4 19 N N N N N X N — — N 13.6 XI L9 15 15 15 1,4 4 24 N N N N N N N — — N 13.9 XI 19 15 15 15 1,4 4 21 N N N N N N N 6 ~30 N 14.4 XI 19 15 15 15 1,4 4 22 N N N X X N N 6 29 N 15.3* XI 19 16 15 15 1,4 \ N 8 6 6 7 12 24 6 6 31 33 15.7 XI 19 15 15 15 1,4 4 13 N N X X X X X 6 24 N 19.0 XI 19 15 15 15 1,4 4 — N N X X X N N 6 N 29 20.4* XI 19 17 15 15 1,4 4 N 8 6 6 7 12 24 6 6 26 26 20.8 XI 20 15 15 N 1,4 4 5 X N N N N N X 6 21 N 22.0* XI 18 17 16 16 1,4 4 N 8 6 6 7 12 24 6 6 34 30 juvenile H. spinosus utilize different areas, perhaps far- ther offshore, as nurseries. Nautichthys oculofasciatus (Girard) (Figure 25; Tables 33, 34) Literature. Larvae of this species were illustrated (7.5, 13 mm SL) and briefly described by Blackburn (1973) and young stages (9.5, 13, 17, 26 mm TL) by Marliave (1975). Distinguishing Features. Larvae hatch at a relatively large size, probably ~7 mm SL. The largest pelagic specimens reported are ~16-17 mm in which all fins are developed except the pelvics. A transformed juvenile, 26 mm TL, was taken in a benthic habitat (Marliave 1975) indicating transformation probably occurs between these sizes. The larvae are distinctively pigmented over the lat- eral surface of the body, except for the caudal peduncle, with melanophores extending onto the dorsal and anal finfolds and fins in the middle of the tail region. Melano- phores also occur over the head, at the base of the pec- toral fin, on the pectoral fin rays as a band near the fin margin, and over the dorsolateral surface of the gut. Pig- ment is added to the spinous dorsal fin as the anterior rays become elongate. Four spines develop on the preopercle appearing as bumps. Enlarged bumps also develop in the parietal region, anterior to it along the parietal ridge, in the postocular region, nasal region, and several in the post- temporal-supracleithral area. None of these appear as pronounced sharp spines. The length of the pelvic fins is striking, 42 and 59% SL in our two specimens. Snout to anus length is relatively long, 56 and 51% SL while body depth at the pectoral fin base is relatively slender, 21 and 23% SL. The number of myomeres (>40) is relatively high for a cottid. The larvae have an agonidlike appearance. Discussion. Only one species of Nautichthys occurs off 40 Table 32. -Measurements (mm) of young of Hemilepidotus spinosus. (Specimens between dashed lines are undergoing notochord flexion.) Body depth 2d preoper- Body Head Snout Eye Snout to at pectoral Pectoral cular Parietal length length length diameter anus length fin base fin length spine length spine length 5.0 0.88 0.12 0.46 1.9 0.82 0.24 — — 6.2 1.1 0.16 0.56 2.6 0.94 0.31 — — 6.6 1.0 0.24 0.54 2.5 0.98 0.35 — — 6.6 1.4 0.24 0.56 2.6 1.0 0.47 — 0.04 7.2 1.4 0.31 0.62 2.8 1.2 0.47 — 0.04 8.3 1.9 0.39 0.82 3.6 1.6 0.47 0.24 0.10 8.6 1.6 0.35 0.78 3.5 1.5 0.66 0.22 0.12 8.9 1.7 0.23 0.82 3.5 1.7 0.74 0.27 0.20 9.1 1.9 0.39 0.90 3.8 1.8 0.78 0.27 0.20 10.1 2.3 0.40 1.2 3.9 2.0 0.82 0.39 0.23 10.6 2.5 0.39 1.2 4.5 2.1 0.82 0.35 0.31 11.0 3.1 0.39 1.3 5.4 2.7 1.5 0.58 0.23 11.0 3.1 0.48 1.3 5.5 3.0 1.5 0.70 -0.27 11.2 3.3 0.40 1.6 5.5 3.0 1.6 0.66 0.31 11.8 3.8 0.70 1.4 6.2 3.4 1.8 0.72 0.24 11.8 3.8 0.80 1.6 6.7 3.4 1.9 0.78 0.31 13.4 4.7 0.96 1.9 7.0 4.2 2.4 0.88 -0.2 13.6 4.6 0.96 2.0 7.4 4.2 2.7 0.88 — 13.9 5.1 1.0 1.8 7.4 4.3 2.9 0.9 — 14.4 5.4 1.0 1.8 8.2 5.0 2.9 BR 1 — 15.7 5.7 1.4 2.0 9.3 5.0 3.4 1.1 — 19.0 7.4 1.7 2.2 11.5 6.4 4.8 1.1 — 20.8 7.8 2.2 2.2 12.5 6.6 5.4 1.1 — 'BR = Broken. Nautichthys oculofasciatus II. 7mm NL Figure 25. — Larvae of Nautichthys oculofasciatus. 41 Oregon and the larvae are striking in appearance. The larvae are rare in our collections. Only two specimens have been taken, both 9 km off Newport, Oreg., in March. Larvae of two additional northeast species N. pribilovius and N. robustus are unknown. Table 33. — Meristics from larvae of Nautichthys oculofasciatus. (Specimen above dashed line is undergoing notochord flexion.) Dorsal Dorsal Anal Pectoral Ventral Pelvic fin Preoper- midline Body length fin spines fin rays fin rays fin rays Left Right spine and rays cular spines melano- phores 11.7 vn 28 19 13 12 - - N 2 16.6 IX 28 19 14 N — 4 N 'Not present as a distinct row. 2 N = Not examined. Radulinus asprellus Gilbert (Figures 26, 27; Tables 35, 36) Literature. Larvae of this species have not been pre- viously described. Distinguishing Features. The smallest larvae we have collected are ~4.5-5 mm long and probably recently hatched. The largest specimens captured pelagically, ~ 14-15 mm, have developed most juvenile character- istics including adult fin complements, increased pigmentation, and lateral line scales. Larvae of R. asprellus are one of the most heavily pigmented among cottids. Melanophores occur over most Table 34. — Measurements (mm) of larvae of Nautichthys oculofasciatus. (Specimen above dashed line is undergoing notochord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper - cular spine length Parietal spine length 11.7 3.4 0.80 1.1 6.6 2.5 4.9 - - 16.6 4.5 1.1 1.3 8.4 3.8 9.8 0.20 — Table 35. — Meristics from young of Radulinus asprellus. (Specimens between dashed lines are undergoing notochord flexion.) [* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; *** = Present but not countable as distinct melanophores; B = Bases only; D = Damaged; N = Not examined.] Pelvic Ventral Caudal fin rays Anal Pectoral fin Preoper- midline** * Dorsal Ventral Vertebrae Branchi- Body Dorsal fin fin fin rays spine cular melano- Second- Prin- Prin- Second- Abdom- Cau- ostegal length Spines Rays rays Left Right &rays spines phores ary cipal cipal ary inal dal rays 4.7 — — — — — NNNNNNNN 5.2 — — — — — — N NNNN NNN 5.9* _____ N N 6.1 NNNNNNNN 6.7 — — — — — — — N NNNN NNN 7.2 — — — — — — 3 N NNNN NNN 7.8* _____ _ N N ____ ___ 7.9 — — — — — — — N NNNN NNN 8.5* _____ N N — - — ___ 9.1 -22B-** 18B — — NNNNNNNN 9.6 VII 21 22 — — 3 N NNNN NNN 10.1 -21-** 22 18N NNNNNNNN 10.9 X 22 22 19 N Buds 4 N NNNN NNN 11.4 -23- ** 22 18 18 Buds 4 N N N N N N N N 11.4* X 21 23 — — — N N — 5 5 — — — — 11.5 IX 22 24 18 18 Buds 4 N X N N N N N N 11.7* X 22 22 16 N — N N — 5 5 — 12 20 6 12.0 IX 22 23 17 17 Buds 4 N N N N N N N N 12.0* X 21 24 18 19 1,3 N N — 6 6 — 12 23 6 12.6 IX 20 22 19 18 1,3 4 X N X X N N N N 12.6 X 23 24 19 19 1,3 4 N X N N N N N N 13.0 IX 22 24 18 19 Buds 4 N N N N X N N N 13.0* vm D 23 18 18 1,3 N N — 6 6 — 12 27 6 13.9* X 22 23 18 18 1,3 N X 3 6 6 2 12 27 6 14.1 IX 23 23 18 L8 1,3 4 N X N N N N N N 14.1 X 22 23 19 19 1,3 4 N N X N N N N N 14.4 IX 22 24 18 18 1,3 4 N N N N N N N N 14.4 X 23 24 19 19 1,3 4 N N X N N N N N 14.7* IX 22 23 19 19 1,3 N N 3 6 6 2 12 27 6 15.2* IX 23 24 18 18 1,3 N N 3 6 6 2 12 27 6 42 Radulinus asprellus 4.7mm NL 7.9mm NL ■ Mas V«+>*' 2**£ s# <*&> 9.6mm NL Figure 26. — Larvae of Radulinus asprellus. of the body including the head, most of the gut, along dorsal and ventral midlines, and laterally except dorso- lateral^ above the body midline over the gut region and dorsally and laterally on the tail tip. This pigment is somewhat darker along the dorsal and ventral body midlines. Some ventral midline melanophores extend beyond the end of the lateral body pigment onto the tail tip, particularly in smaller specimens. A distinctive series of elongate melanophores occurs along the lateral midline appearing almost as a solid line. The presence of these melanophores help distinguish larvae of R. asprel- lus from Scorpaenichthys marmoratus which they resem- ble at small sizes. Pigmentation generally increases with development with only the tail tip or caudal peduncle re- maining unpigmented. The lateral midline pigment is noticeable on all but the largest specimens. Four small spines develop along the preopercular margin, visible by ~ 10-11 mm. They never become prominent. A parietal and nuchal spine develop as 2 minute spines which soon appear only as bumps. A nasal spine is evident on the largest transforming specimens ~14-15 mm. The gut is rather distinctively coiled with snout to anus length ranging from ~44 to 54% SL. The snout becomes quite pointed further helping to distinguish these larvae from S. marmoratus. The larvae are rela- tively slender with body depth at the pectoral fin base never exceeding 25% SL during the pelagic phase and always <20% SL in specimens <10 mm. The adult com- plement of all fin rays or ray elements is attained by ~11- 12 mm. Discussion. Three species of Radulinus occur in the northeast Pacific. Larvae of R. boleoides are described in 43 Radulinus asprellus ,.y I0.9mm NL 12.6mm SL 14.4mm SL Figure 27. — Young of Radulinus asprellus. the next section. They are distinguishable from R. asprellus on the basis of body proportions and small pig- ment differences as given in that section, at least for the size range described. Larvae of the third species, R. vinculus, are unknown. Larvae of R. asprellus are one of the more heavily pigmented cottids along with, e.g., Scorpaenichyths marmoratus, Rhamphocottus richardsoni, and Blepsias cirrhosus (Blackburn 1973; Marliave 1975), all of which have melanophores covering most of the body except the tail tip or caudal peduncle. Interestingly only one of these, S. marmoratus, is known to be common in the neuston, indicating larval pigmentation may not reflect habitat preference, at least in these species. Radulinus boleoides Gilbert (Figure 28; Tables 37, 38) Literature. The larvae of this species have not been previously described. Distinguishing Features. We have identified only two larvae as R. boleoides in our collections, 7.5 and 8.7 mm long. At comparative sizes they are somewhat more ad- vanced in development than larvae of R. asprellus. The pigment pattern closely resembles that of R. asprellus with melanophores covering most of the body laterally except the tail tip. A few ventral midline mela- 44 Table 36. — Measurements (mm) of young of Radulinus asprellus. (Specimens between dashed lines are undergoing notochord flexion.) Body depth 2d preoper- Body Head Snout Eye Snout to at pectoral Pectoral cular Parietal length length length diameter anus length fin base fin length spine length spine length 4.7 1.1 0.10 0.51 2.6 0.86 0.47 — — 5.2 1.2 0.20 0.47 2.4 0.82 0.27 — — 6.1 1.3 0.32 0.54 2.8 0.98 0.27 — — 6.7 1.6 0.35 0.62 3.3 1.1 0.35 — — 7.2 1.9 0.43 0.58 3.4 1.2 0.35 — — 7.9 1.8 0.39 0.70 3.5 1.3 0.55 — — 9.1 2.3 0.56 0.56 4.2 1.6 0.55 — — 9.6 2.7 0.82 0.82 4.6 1.8 0.78 — — 10.1 2.4 -0.50 0.88 4.9 1.8 0.80 — — 10.9 2.9 0.78 1.0 5.9 2.6 1.1 — — 11.4 3.0 0.72 1.0 6.0 2.6 1.8 — — 11.5 3.5 0.88 1.0 6.2 2.5 2.0 0.27 — 12.0 3.6 1.0 1.2 6.3 2.5 2.1 0.23 — 12.6 4.1 0.88 1.4 6.6 3.0 3.1 0.35 — 12.6 3.7 0.88 1.21 6.6 3.0 l.H 0.23 — 13.0 3.6 0.96 1.1 6.2 2.6 1.7 0.27 — 14.1 4.5 0.96 1.4 7.6 3.5 2.0 0.40 — 14.1 4.8 1.3 1.4 7.0 2.8 2.7 0.46 — 14.4 4.7 1.1 1.5 7.0 3.0 3.7 0.43 — 14.4 4.2 1.0 1.4 7.0 3.4 2.2 0.35 — Radulinus boleoides 'jfJjA. &C-SJ 8.7mm NL Figure 28. — Larva of Radulinus boleoides. Table 37. -Meristics from larvae of Radulinus boleoides. mens are undergoing notochord flexion.) (Speci- Ventral Dorsal Dorsal Anal Pectoral Pelvic fin Preoper- midline Body fin fin fin fin rays spine cular melano- length spines rays rays Left Right and rays spines phores 7.5 VIII 20 15 18 N 2 Buds — N 8.7 IX 16 20 19 N 1,3 — N 'Present but not countable as distinct melanophores. 2 N = Not examined. nophores extend onto the tail tip. They differ somewhat in the posterior extent of the lateral pigment which is less in R. boleoides than in R. asprellus of similar size, i.e., a greater portion of the tail tip is pigmentless in R. boleoides. Larvae of R. boleoides also have a distinctive series of melanophores along the lateral midline as do R. asprellus. No head spines are visible on the two flexion larvae in our collection, although they may develop later. Table 38. — Measurements (mm) of larvae of Radulinus boleoides. (Specimens are undergoing notochord flexion.) Body Head length length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 7.5 2.4 8.7 2.6 0.58 0.64 0.70 0.78 4.4 5.0 1.7 2.0 0.98 1.2 — — 45 The coiled gut is rather prominent and distinctive, as in R. asprellus, but snout to anus length is considerably greater in R. boleoides (58% SL) than R. asprellus (44- 47% SL) of similar size. Body depth at the pectoral fin base is also greater in R. boleoides (22-23% SL) than R. asprellus (16-18% SL). The pectoral fin is relatively longer in R. boleoides (13-14% SL) compared to R. asprellus at that size (5-7% SL). Both specimens of R. boleoides are undergoing notochord flexion and adult fin ray complements have not been attained, explaining the discrepancies in the fin ray counts in Table 37. Discussion. Identification was based on the strong sim- ilarity of these larvae to R. asprellus and the major dif- ferences in body proportions which were obviously not caused by distortion. The only other species of Radu- linus which occurs off Oregon is R. boleoides and it is relatively rare, explaining the paucity of larvae in our collections. Our two specimens were taken 9 and 18 km off the Oregon coast in March and April. Rhamphocottus richardsoni Gunther (Figure 29; Tables 39, 40) Literature. Larvae (6.7, 10 mm SL) were illustrated and briefly described by Blackburn (1973) and young (10, 11.5, 15 mm TL) by Marliave (1975). Distinguishing Features. Hatching occurs ~6-7 mm and transformation at ~ 14-15 mm (Blackburn 1973; Marliave 1975). Pelagic specimens in our collections range from 6.0 to 11.8 mm. Larvae are rather heavily pigmented with melano- phores covering most of the body except the tail tip or caudal peduncle. Pigment covers the cheeks, and ventral surface of the gut on the smallest specimens. Pigment does not appear noticeably darker along the dorsal and ventral body margins and no distinct line of melano- phores occurs along the lateral midline. Some melano- phores occur on the ventral finfold and occasionally on the dorsal finfold. The preanal finfold is darkly pigmented. Melanophores are present on the base of the pectoral fin. The caudal peduncle remains unpigmented through the pelagic period. Body pigment changes little with development except as transformation approaches, pigment is added to the spinous dorsal fin and pecto- ral fin base, and pigment bands begin to show on the body. Only 1 spine develops on the dorsal margin of pre- opercle, evident by ~10 mm. Smaller spiny projections appear along the preopercular margin which are modi- fied scales developing over the head region. A postocular spine is apparent over the eye by ~8 mm and becomes rather pronounced in larger specimens. A parietal and a nuchal spine develop and fuse together appearing as one. Two spines become obvious in the posttemporal-supra- cleithral region and a cleithral spine and nasal spine are evident on our largest specimens. Body shape is characteristic. Larvae are deep bodied, increasing from 29 to 40% SL between 8 and 12 mm. Snout to anus distance is >60% SL. By 10 mm, larvae begin to resemble adults in form. The larvae have a dis- tinct pigmented preanal finfold which remains evident anterior to the anal opening at least until ~12 mm. To our knowledge, a preanal finfold has been reported in Scorpaenichthys marmoratus (O'Connell 1953), Hemi- tripterus americanus (Warfel and Merriman 1944; Fui- man 1976), possibly H. villosus as indicated on a figure by Okiyama and Sando (1976), Myoxocephalus quadri- cornis, M. scorpius (Khan 1972), possibly M. aenaeus based on figures by Lund and Marcy (1975), possibly Gymnocanthus tricuspis and G. herzensteini based on illustrations by Rass (1949) and Kyushin (1970), Eno- Table 39.— Meristics from larvae of Rhamphocottus richardsoni. (Specimen above dashed line is undergoing notochord flexion.) (* = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; *** = Present but not countable as distinct melanophores; B = Bases only; N = Not examined.] Dorsal fin Spines Rays Anal fin rays Pectoral fin rays Left Right Pelvic fin spine & rays Preoper- cular spines Ventral midline melano- phores Caudal fin rays Vertet Abdom- inal rae Cau- dal >** Dorsal Second- Prin- ary cipal Ventral Branchi- Body length Prin- cipal Second- ary ostegal rays 8.4 -14B-** 6B - - Buds - N \ N N N N N N 10.6 11.7 11.8* VIII 13 VIII 12 VIII 13 6 6 7 16 15 16 16 16 16 1,3 1,3 1,3 1 1 1 N N N N N 1 N N 6 N N 5 N N 1 N N 12 N N 15 N N 6 Table 40. — Measurements (mm) of larvae of R hamphocottus richardsoni. notochord flexion.) (Specimen above dashed line is undergoing Body length Head length 2.6 Snout length 0.55 Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 8.4 0.94 5.5 2.4 1.0 - — 10.6 11.7 4.1 4.8 1.0 1.0 1.1 1.3 7.7 8.1 4.0 4.7 2.2 2.6 — 0.43 0.35 46 Rhamphocottus richardsoni £2zr~/ r™ •>3«^*W 3V»*' l S.*irV3*>.~ \" ' /'k-~- 1 ** * *'■*' • ' < ; @ 8.4mm NL mj$$™- : - 10.6mm SL II. 7mm SL Figure 29. — Young of Rhamphocottus richardsoni. 47 phrys bison (Misitano 1978), and E. bubalis and possibly E. lilljeborgi (Russell 1976), within the Cottidae. Discussion. Rhamphocottus is a monotypic genus. The larvae are distinctive. They share the character of having a rather heavily pigmented body, except for the tail tip, with a number of other northeast Pacific species, as dis- cussed under Radulinus asprellus, but their character- istic body shape renders them outstanding except perhaps at the smallest sizes. The heavily pigmented preanal finfold, and presence of pigment on the anal fin- fold, base of the pectoral fin, and the ventral surface of the gut help distinguish small larvae. The body becomes covered with distinctive spinelike scales while still in the pelagic habitat. The larvae are relatively rare in our col- lections with only seven identified specimens, all taken 2-9 km off the Oregon coast in March and April. Scorpaenichthys marmoratus (Ayres) (Figures 30, 31; Tables 41, 42) Literature. Eggs and young (5.85, 6.26, 10, 17, 30, 48 mm) of this species were described by O'Connell (1953). Distinguishing Features. Larvae are between 4 and 6 mm long at hatching. Most of our pelagic specimens are <15 mm. By ~13-14 mm they have a juvenile appear- ance and fully formed fins. However, we have captured a 21.6 mm specimen pelagically and a 35 mm specimen in a neuston tow 9 km off Newport, Oreg. O'Connell (1953) reported specimens as large as 29 mm were collected with dip nets. Larvae are very heavily pigmented over most of the body, usually appearing darker than larvae of Radulinus Scorpaenichthys marmoratus 5.3mm NL 7.5mm NL 8.6mm NL Figure 30. — Larvae of Scorpaenichthys marmoratus. 48 Scorpaenichthys marmoratus 10. 4mm SL 13.8mm SL Figure 31. — Young of Scorpaenichthys marmoratus. asprellus or Rhamphocottus richardsoni. In small larvae melanophores cover the head, the gut except the ventral margin, the dorsal body margin, the ventral margin of the tail, and the lateral body surface except for the tail tip. The lateral pigment extends more posteriorly than in Radulinus spp. The dorsal body margin, dorsal surface of the gut, and postanal ventral body margin appear darker than the rest of the body. The pectoral fin, preanal fin- fold, and body finfold are unpigmented. With develop- ment, melanophores are added on the head and laterally except along the ventral margin of the gut which remains unpigmented until ~8-9 mm and the caudal peduncle which retains an unpigmented band at least until 14-15 mm. As the median fins develop the dorsal and ventral body margins do not appear as heavily outlined. On specimens ~13-14 mm a discrete melanophore can be seen at the base of most dorsal fin soft rays. Larvae gen- erally do not develop a distinct line of pigment along the lateral midline as observed in larvae of Radulinus spp. The margin of the preopercle develops 4 spines, not 3 as reported by O'Connell (1953), but only the upper 3 spines become pronounced. They are first noticeable ~8 49 mm, and become striking in postflexion stages with the length of the 2d spine ranging from 22 to 26% HL. Raised bumps develop in the parietal region, the posttemporal- supracleithral region, and the cleithral region, but spines never form. A nasal spine becomes obvious by ~ 13-14 mm. Preflexion larvae are relatively slender with body depth at pectoral fin base ranging from 17 to 19% SL. Table 41. — Meristics from young of Scorpaenichthys marmoratus. (Specimens between dashed lines are undergoing notochord flexion.) [* = Specimens stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; *** = Present but not countable as distinct melanophores; B = bases only; N = Not examined.] Pelvic Ventral Caudal fin rays Dorsal fin Spines Rays Anal fin rays Pectoral fin rays fin spine & rays Preoper- cular spines midline* melano- phores ** Dorsal Ventral Vertebrae Abdom- Cau- inal dal Branchi- Body Second- ary Prin- cipal Prin- cipal Second- ary ostegal length Left Right rays 5.1* N N — — — — — — — 5.3 X N N N N N N N 5.4 X N N N N N N N 5.8 N N N N N N N N 6.1 N N N N N N N N 6.3 N N N N N N N N 6.6* N N 7.5 N N N N N N N N 7.5* N X 7.7* N N 8.1 2 N N N N N N N N 8.1 -15B-** 12B — — — 4 N N N N N N N N 8.2* N N 8.6 -16B-** 13B — — — 2 N N N N X N N N 8.7 -15- ** 13 11 14 — 4 N N N N N N N N 9.1* 12 10 N N — 6 6 — 14 16 6 9.3 -16- ** 13 14 15 Buds 4 N N N N X N N N 10.1* X 18 14 13 13 — N N 5 6 6 2 14 19 6 10.4 XI 17 12 15 15 Buds 4 N N N N N N N N 10.4 XI 15 11 14 14 Buds 4 N N N X X X N N 10.7 XI 17 12 15 15 Buds 4 N N N N N X N N 10.9* XI 17 14 15 15 1,3 N N 7 6 6 6 15 21 6 11.5 X 16 12 14 15 1,5 4 N N X X X N N N 12.2* XI 17 14 16 16 1,5 N N 9 6 6 5 15 21 6 13.8 XI 17 12 15 15 1,5 4 N N X N X X N N 13.8* XI 17 13 N 15 1,5 N N 9 6 6 9 15 21 6 14.7 XI 15 1.3 15 15 1,5 4 X N N N X N N N 21.6 X 16 13 15 15 1,5 4 N X N N N N N N Table 42. — Measurements (mm; Body length of young of Scorpaenichthys marmoratus. undergoing notochord flexion.) (Specimens between dashed lines are Body depth 2d preoper- Head Snout Eye Snout to at pectoral Pectoral cular Parietal length length diameter anus length fin base fin length spine length spine length 5.3 1.1 0.27 0.44 2.8 0.90 0.44 — — 5.4 1.2 0.27 0.56 2.9 0.98 0.40 — — 5.8 1.3 0.26 0.56 3.1 0.98 0.43 — — 6.1 1.4 0.30 0.56 3.1 1.1 0.48 — — 6.3 1.3 0.36 0.56 3.1 1.2 0.46 — — 7.5 1.6 0.27 0.66 3.7 1.2 0.82 — — 8.1 1.9 0.31 0.82 4.6 1.9 0.55 — — 8.1 2.5 0.39 1.05 4.6 2.1 0.82 0.32 — 8.6 1.9 0.35 0.68 4.2 1.6 0.70 — — 8.7 2.2 0.39 0.98 5.2 2.4 0.96 0.42 — 9.3 3.1 0.40 1.4 5.8 3.2 1.6 0.70 — 10.4 3.8 0.72 2.2 6.2 3.3 1.8 0.90 — 10.4 3.1 0.48 1.4 5.9 3.3 1.8 0.78 — 10.7 3.4 0.48 1.7 6.5 3.8 2.0 0.78 — 11.5 3.8 0.40 1.8 7.4 3.8 2.2 1.01 Bump 13.8 5.4 0.96 2.2 8.9 4.6 3.0 1.24 Bump 14.7 5.4 0.88 2.5 9.7 5.2 3.4 1.36 — 21.6 8.8 1.92 2.7 15.5 8.2 6.2 1.44 — 50 They become much deeper bodied with development with a body depth of 32-35% SL in postflexion stages. Snout to anus length is relatively long, increasing from 49-53% SL in preflexion larvae to 57-66% SL in post- flexion larvae. The snout develops a rounded appear- ance, never pointed as in Radulinus sp. An unpigmented preanal finfold remains evident up to ~13-14 mm. The presence of such a preanal finfold has been reported for relatively few cottid genera (see Discussion under Rham- phocottus richardsoni). Discussion. Scorpaenichthys is a monotypic genus. The larvae are rather distinctive and one of the most heavily pigmented forms of northeast Pacific cottid species. Small larvae appear somewhat similar to Radulinus spp. but have lateral pigment extending further toward the tail tip, no ventral midline melanophores posterior to the lateral pigment and usually no distinct line of melano- phores along the lateral midline. Occasionally newly hatched larvae will have very few melanophores on the lateral body surface causing them to resemble small Hemilepidotus spinosus. However, their gut is longer and the dorsal and ventral midline pigment does not extend as far onto the tail tip as in H. spinosus. Larvae of S. marmoratus appear to be strongly asso- ciated with the neuston. O'Connell (1953) reported dip net collections contained specimens 7.4-29 mm long, and we have taken specimens 4.0-35 mm in neuston tows off Oregon. One 10-min neuston tow taken 9 km off New- port, Oreg., in March contained 829 larvae, mostly 4-5 mm long and recently hatched. Richardson and Pearcy (1977) reported them to be a coastal form, as the larvae were only taken within 28 km of the coast in oblique plankton tows. However, we have since collected them in the neuston as far as 65 km offshore, well beyond the continental shelf-slope break. Cottoid Type A (Figure 32; Tables 43, 44) Literature. No published descriptions of this form exist although illustrations were given by Richardson and Bond 9 as "Globby Type A." These larvae were called 'Richardson, S. L., and C. E. Bond. 1978. Two unusual cottoid fishes from the northeast Pacific. Unpubl. manuscr., 6 p. Gulf Coast Research Laboratory, Ocean Springs, MS 39564, or Department of Fish- eries and Wildlife, Oregon State University, Corvallis, OR 97331. Psychrolutes-like sp. 1 by Richardson (footnote 4) and Richardson and Pearcy (1977). Distinguishing Features. Three larval specimens (9.8, 12.8, 13.4 mm) of this unusual looking form have been collected off Oregon. The smallest specimen is under- going notochord flexion and the two larger specimens are postflexion larvae. The largest larva, which was stained, has the full complement of dorsal, anal, pectoral, and pelvic fin rays but secondary caudal rays are still developing and the vertebral centra are not completely ossified, thus transformation occurs at a larger size. The larvae are pigmented over the entire body except at the tail tip. The basal portion of the pectoral fin is pigmented, but its distal portion and the dorsal and anal finfolds are unpigmented. No spines are apparent in the head region. The belly is covered with small, thumb-tack prickles which extend dorsolateral^ in lesser numbers and a few are on the tail region in the larger specimens. These larvae are globose in shape, unlike any other known cottid larvae. The outer skin is loose. They are deep bodied with depth at the pectoral fin base 42 and 56% SL in the two specimens illustrated. Snout to anus distance is also great, 58 and 79% SL, respectively. The pelvic fins appear to be inserted in pockets in the skin with only the tips exposed in all three larvae. This condi- tion may be variable as reported in adults in a Southern Hemisphere cottoid genus Neophrynichthys (Nelson 197- 7) and as observed in larvae of a Bering Sea cottoid form (unpubl. data) referred to as Malacocottus zonurus - Type 1 by Richardson and Bond (footnote 9). The first dorsal fin appears to be embedded in the skin and the second dorsal and anal fins are partially skin covered. Discussion. Meristics, particularly the high number of pectoral fin rays and the possession of seven branchi- ostegal rays, agree with only one species described from the area, Psychrolutes phrictus (Stein and Bond 1978). Young stages of P. phrictus <50 mm also have prickles in the belly region similar to these larvae. Based on this information alone our Cottoid Type A larvae would ap- pear to be P. phrictus. However, discovery of a single 26.4 mm juvenile (Richardson and Bond footnote 9) which also agrees with P. phrictus in meristics and possession of belly prickles has lead to uncertainty regarding its iden- tity. This unidentified juvenile differs drastically in body form from a 35 mm P. phrictus, primarily in relative Table 43. — Meristics from larvae of Cottoid Type A. (Specimen above dashed line is undergoing notochord flexion.) = Specimen stained with Alizarin Red S; ** = Dorsal fin spines and rays not distinguishable; D = Damaged; N = Not examined.] Dorsal fin Spines Rays Anal fin rays Pectoral fin rays Left Right Pelvic fin spine & rays Preoper- cular spines Ventral midline melano- phores Caudal Dorsal Second- Prin- ary cipal fin rays Ventral Vertebrae Abdom- Cau- inal dal Branchi- Body length Prin- cipal Second- ary ostegal rays 9.8 -15-** 11 23 23 - - - N N N N N N N 12.8 13.4* -16-** VIII 20 11 14 24 26 D 26 1,3 - - N N 6 N 7 N 4 N 12 N 23 N 7 51 Cottoid Type A 9.8 mm SL 12.8 mm SL Figure 32. — Larvae of Cottoid Type A. Arrows indicate the position of the anus. Table 44. — Measurements (mm) of larvae of Cottoid Type A. (Specimen above dashed line is undergoing notoehord flexion.) Body length Head length Snout length Eye diameter Snout to anus length Body depth at pectoral fin base Pectoral fin length 2d preoper- cular spine length Parietal spine length 9.8 3.6 0.66 1.2 5.7 4.1 1.7 - - L2.8 6.1 1.5 1.7 10.1 7.2 3.7 — — 52 body depth (51% and 31% SL, respectively) and snout to anus length (61% and 51% SL, respectively). The unidentified 26.4 mm juvenile agrees much more closely with our Cottoid Type A larvae than with the 35 mm P. phrictus juvenile. It seems unlikely that a drastic change in body proportion would occur during the juvenile period between 26 and 35 mm. And the 26.4 mm speci- men does not appear to be distorted, based on radio- graphs. It also seems unlikely that two such unusual forms would exist in the northeast Pacific that agree so well in meristics and belly prickles. However, the identi- fication of our Cottoid Type A larvae as P. phrictus can- not be positively made until additional material becomes available to provide a conclusive linkage between our lar- vae and identified juveniles. If our Cottoid Type A larvae are found to be P. phrictus, then placement of this species in the genus Psy- chrolutes is obviously in error based on the extreme dis- similarity of these larvae and those of P. paradoxus de- scribed by Blackburn (1973) (10.3 mm SL) and Marliave (1975) (10.5, 13, 14 mm TL), and its generic position should be reexamined. Alternatively, these larvae could represent a new species and possibly a new genus of cot- toid fish from the northeast Pacific. The term "cottoid" has been used here because of the dissimilarity of these larvae to any known larvae in the Family Cottidae. NOTES ON OTHER NORTHEAST PACIFIC COTTID LARVAE Larvae are known for 14 additional species of north- east Pacific cottids not described in this paper (Table 45). Those in the genera Artedius, Clinocottus, and Hemilepidotus (including Melletes from the Bering Sea) were discussed under the descriptions of related species. Larvae of Oligocottus spp. were discussed under Cottidae Type 3. Comments on the larval characters in remaining genera, Blepsias, Dasycottus, Gilbertidia, ?Malaco- cottus, Myoxocephalus, Orthonopias, and Psychrolutes are included here for comparative purposes, based on the literature cited in Table 45 and personal observations of specimens not in the Oregon State University Larval Fish Reference Collection. Larvae of Blepsias cirrhosus are one of the dark, heavily pigmented forms along with Radulinus spp., Rhamphocottus richardsoni, and Scorpaenichthys marmoratus. The entire body is covered with dense melanophores except for the tail tip, the ventral surface of the gut, and the cheek. The pectoral fins are un- pigmented and no melanophores occur on the finfold as in R. richardsoni. They have no preanal finfold as in R. richardsoni and S. marmoratus. They apparently hatch at a relatively large size. The smallest larvae reported from plankton collections are 10 mm TL and just be- ginning to undergo notochord flexion. Thus, they are generally less developed than the above-mentioned dark cottid larvae at comparable sizes. The larvae appear more slender with respect to body length than S. marmo- ratus and R. richardsoni and relative snout to anus length is less than in R. richardsoni. They do not develop the pronounced preopercular spines seen in S. mar- moratus. The largest specimen reported from plankton collections is 25.5 mm TL and is apparently trans- formed, or nearly so. Table 45.— Additional northeast Pacific cottid species for which larvae-and young are known. Species Artedius lateralis Blepsias cirrhosus Clinocottus analis Clinocottus recalvus Dasycottus setiger Gilbertidia sigalutes Hem ilepidotus jordani Hemilepidotus zapus ?Malacocottus zonurus - Type 1 Myoxocephalus polya- canthocephalus Oligocottus maculosus Oligocottus snyderi Orthonopias triads Psychrolutes paradoxus Reference Budd 1940 Marliave 1975 Blackburn 1973 Marliave 1975 Eigenmann 1892 Budd 1940 Morris 1951 Blackburn 1973 Blackburn 1973 Marliave 1975 Gorbunova 1964 Peden 1964 Peden 1964 Richardson and Bond 1978 Richardson (unpubl. data) Blackburn 1973 Stein 1972 Stein 1973 Stein 1972 Bolin 1941 Blackburn 1973 Marliave 1975 Size illustrated 4.1mmSL 4, 8, 11, 14 mm TL 12.2 mm SL 10, 14, 19, 25.5 mm TL ~4 mm ~4 mm 4.6,5.0,7.6,8.3,9.9, 10.8,18.0, 24.3 mm TL 7.4 mm SL 7.3, 9.5 mm SL 7, 13,15, 25, 34 mm TL 6.4, 10.7, 13.0 mm ~20mm ~20mm 7.0,9.8, 14.2, 24.0 mm SL 6.6, 7.0, 8.8, 9.8, 10.4, 14.2, 24.0 mm SL 7.7, 10.7 mm SL 4.6-5.2,6.0, 6.6, 9.2 mm TL 4.6-5.2, 6.0, 6.6, 9.2, 12.2 mm TL 4.5-4.75, 5.5 mm TL ~3-4 mm SL 10.3 mm SL 10.5, 13, 14, 13 mm TL See text footnote 9. 53 Larvae of Dasycottus setiger are characterized by a relative lack of pigment on the body except on the head and over the relatively short gut. Melanophores in these two regions are low in numbers but are large in appear- ance, thus covering most of the head and abdominal cavity. Pigment is absent from the postanal ventral midline. Pigment is present on the underside of the pec- toral fin and later is added to the blade and outer side of the fin. The general pigment pattern, except for the pigmented pectorals, resembles our Triglops sp. but the number of myomeres is much lower for D. setiger, ~34- 35. The smallest larvae reported are 7.4 mm and pre- sumed to be recently hatched. Fin rays are formed by 12 mm and the pigment pattern remains unchanged at least until 19 mm. A 24 mm SL specimen observed by us has begun to acquire juvenile pigmentation. Small larvae of Gilbertidia sigalutes and Psychrolutes paradoxus are similar to each other. Both hatch at ~6-7 mm. The smallest larvae resemble those of liparids in general shape. Both have moderate melanistic pigment over the head, nape, gut, and on the pectoral fins. Post- anal ventral midline melanophores are absent. On both, pigment is added to the lateral body surface above the abdominal cavity and posterior to it, with relatively more pigment present in P. paradoxus than G. sigalutes of similar size particularly in larger specimens. Both develop the appearance of having loose skin. The number of pectoral fin rays separate the two with > 19 for P. para- doxus and <18 for G. sigalutes. These are usually count- able on all but the smallest larvae. Additional definitive differences between the smallest larvae of these two spe- cies, based on plankton preserved material, remain to be described. Blackburn (1973) indicated differences exist in the shape of melanophores on the pectoral fin and head, but we have not found this to be consistent and reliable. Marliave (1975) stated that freshly preserved larvae of P. paradoxus have more melanin in the pectoral fins which obscures their orange color while the pectoral fins of G. sigalutes appear bright orange. Thus there may be real differences in the melanistic pigment patterns on the pectoral fins of small larvae, but the limits of varia- tion of those patterns remain to be defined. Pectoral fin pigment differences are apparent in larger preserved specimens with those of G. sigalutes being much darker and having unpigmented circles near the margin. Marliave also stated that P. paradoxus larvae develop a notch dorsally in the caudal finfold during fin ray forma- tion whereas G. sigalutes larvae do not. This was not ob- vious on all specimens we have examined. Marliave observed that P. paradoxus settles from the plankton ~13-14 mm TL while G. sigalutes remains pelagic until >20 mm TL. Larvae of an additional form from the Gulf of Alaska and Bering Sea, tentatively called Malaco- cottus zonurus - Type 1 by Richardson and Bond (foot- note 9), are also very similar in appearance to larvae of G. sigalutes and P. paradoxus. Small larvae of all three are nearly identical but larvae of M. zonurus - Type 1 develop 4 preopercular spines with an anteriorly di- rected accessory spine at the base of the second spine. This secondary spine is visible on larvae by ~7 mm upon close examination. Pectoral fin ray counts, ~20-22, will further separate it from G. sigalutes. Often the pelvic fins of M. zonurus - Type 1 appear to be withdrawn into pockets in the skin, although this condition is variable. This form has very loose skin and remains pelagic at least until ~24 mm. Positive identification of these larvae as M. zonurus awaits resolution of apparent taxonomic problems in the genus Malacocottus (Howe and Rich- ardson footnote 3; Richardson and Bond footnote 9). In larvae of Myoxocephalus polyacanthocephalus, which hatch ~7 mm, melanophores cover most of the anterior one-half to two-thirds of the body except the ventral surface of the abdomen. The posterior one-third of the body remains conspicuously unpigmented except for a series of small melanophores along the ventral midline which extends to near the tail tip. The pigment pattern is similar to other dark cottid larvae, e.g., Radu- linus asprellus, Rhamphocottus richardsoni, Scorpaenichthys marmoratus, Blepsias cirrhosus, except that a greater proportion of the tail region remains un- pigmented laterally and the unpigmented tail region also retains a series of ventral midline melanophores. The head develops prominent spines including at least 4 pre- opercular, a parietal, a nuchal, a postocular, and several in the posttemporal-supracleithral region. Early larvae of Orthonopias triads resemble those of Artedius spp. with which it is closely related (Howe and Richardson footnote 3). At hatching they have pigment over the gut and a series of ~35 postanal ventral midline melanophores. Soon after hatching pigment is added to the interorbital region and on the nape. Apparently no gut diverticula are present. Characters given in the liter- ature are inadequate to distinguish them from Artedius harringtoni except for possibly a higher number of ven- tral midline melanophores. ACKNOWLEDGMENTS Examination of larval cottids from other areas of the northeast Pacific was helpful in working out identifica- tions of our Oregon cottid larvae. We thank the following people with collections from particular areas: northern California — David W. Rice (Lawrence Livermore Labo- ratory); Washington and Puget Sound — Jean R. Dunn and Kenneth D. Waldron (Northwest and Alaska Fish- eries Center, National Marine Fisheries Service, NOAA), Lawrence Moulton (University of Washington), James Blackburn (Alaska Department of Fish and Game formerly of University of Washington), Andrew Lamb (Pacific Environment Institute, Vancouver), British Col- umbia — W. E. Barraclough (Fisheries Research Board of Canada, Nanaimo) and Daniel J. Faber (Canadian Oceanographic Identification Center); Bering Sea and Gulf of Alaska — Pat Wagner and George Mueller (Uni- versity of Alaska). Jean R. Dunn provided reared larvae of Hemilepidotus hemilepidotus. Kevin Howe, Oregon State University, provided transforming and juvenile cottids collected in Oregon tidepools and general infor- mation on tidepool collections. James Blackburn and Jeffrey B. Marliave, Vancouver Public Aquarium, pro- 54 vided specimens of Gilbertidia sigalutes and Psychro- lutes paradoxus for comparative examination. Many people helped in one way or another in working out larval cottid identification problems particularly E. H. Ahl- strom, NOAA, NMFS Southwest Fisheries Center; James Blackburn; Carl E. Bond, Oregon State Uni- versity; Jean R. Dunn; Kevin Howe; Joanne L. Laroche, Oregon State University; Wayne L. Laroche, Oregon State University; Kenneth Waldron; and Sharon Roe, formerly of Oregon State University. We thank E. H. Ahlstrom and Carl E. Bond for reviewing the manu- script and offering helpful comments. Special thanks are extended to Jean R. Dunn whose efforts made this paper possible. This research was supported in part by NOAA- NMFS Contract No. 03-78-M02-120, 1 January 1978 to 30 September 1978. LITERATURE CITED AHLSTROM, E. H., J. L. BUTLER, and B. Y. SUMIDA. 1976. Pelagic stromateoid fishes (Pisces, Perciformes) of the east- ern Pacific: Kinds, distributions, and early life histories and observations on five of these from the northwest Atlantic. Bull. Mar. Sci. 26:285-402. BEARDSLEY, A. J., and C. E. BOND. 1970. Field guide to common marine and bay fishes of Ore- gon. Oreg. State Univ. Agric. Exp. Stn. Bull. 607, 27 p. BLACKBURN, J. E. 1973. A survey of the abundance, distribution, and factors affecting distribution of ichthyoplankton in Skagit Bay. M.S. Thesis., Univ. Washington, Seattle, 136 p. BOLIN, R. L. 1941. Embryonic and early larval stages of the cottid fish Ortho- nopias triads Starks and Mann. Stanford Ichthyol. Bull. 2:73- 82. 1944. A review of the marine cottid fishes of California. Stanford Ichthyol. Bull. 3:1-135. BREDER, C. M., Jr., arid D. E. ROSEN. 1966. Modes of reproduction in fishes. Natural History Press, Garden City, N.Y., 941 p. BUDD, P. L. 1940. Development of the eggs and early larvae of six California fishes. Calif. Dep. Fish Game, Fish. Bull. 56:1-50. EHRENBAUM, E. 1905-1909. Eier und Larven von Fischen. In Nordisches Plank- ton; Zoologischer Teil Erster Band, p. 1-414. Kiel und Leip- zig. EIGENMANN, C. H. 1892. The fishes of San Diego, California. Proc. U.S. Natl. Mus. 15:123-178. ELDRIDGE, M. B. 1970. Larval fish survey of Humboldt Bay. M.S. Thesis, Hum- boldt State Coll., Areata, Calif., 52 p. FEDOROV, V. V. 1973. AlistofBeringSeafish. [In Russ.] Proc. TINRO 87:42-71. [Engl, transl. by Transl. Bur., Multilingual Serv. Div., Dep. Sec. State Canada, Fish. Mar. Ser. Transl. Ser. No. 3271.] FOLLETT, W. I. 1952. Annotated list of fishes obtained by the California Academy of Sciences during six cruises of the U.S.S. Mulberry, conducted by the United States Navy off central California in 1949 and 1950. Proc. Calif. Acad. Sci., Ser. 4, 27:399-432. FUIMAN, L. A. 1976. Notes on the early development of the sea raven, Hemi- tripterus americanus. Fish. Bull., U.S. 74:467-470. GORBUNOVA, N. N. 1964. Breeding and development of hemilipidotine sculpins (Cot- tidae, Pisces). In T. S. Rass (editor), Fishes of the Pacific and In- dian Oceans. Biology and distribution, p. 249-266. Israel Program Sci. Transl. No. 1411. HART, J. L. 1973. Pacific fishes of Canada. Fish. Res. Board Can. Bull. 180, 740 p. HATTORI, S. 1964. Studies on fish larvae in the Kuroshio and adjacent waters. [In Jpn., Engl, synop.j Bull. Tokai Reg. Fish. Res. Lab. 40, 158 p. JONES, A. C. 1962. The biology of the euryhaline fish Leptocottus armatus ar- matus Girard (Cottidae). Univ. Calif. Publ. Zool. 67:321-368. KHAN, N. Y. 1972. Comparative morphology and ecology of the pelagic larvae of nine Cottidae (Pisces) of the northwest Atlantic and St. Lawrence drainage. Ph.D. Thesis., Univ. Ottawa, Ottawa, 234 p. KOEFOED, E. 1907. Croisiere oceanographique dans la mer du Gronland en 1905. Charles Bulens, Brussels, p. 485-500. (Cited by Khan 1972; not seen by authors.) KYUSHIN, K. 1970. Embryonic development and larvae of Gymnocanthus herzensteini Jordan and Starks. Jpn. J. Ichthyol. 17:74-79. LUND, W. A., Jr., and B. C. MARCY, Jr. 1975. Early development of the grubby, Myoxocephalus aenaeus (Mitchill). Biol. Bull. (Woods Hole) 149:373-383. MARLIAVE, J. B. 1975. The behavioral transformation from the planktonic larval stage of some marine fishes reared in the laboratory. Ph.D. Thesis, Univ. British Columbia, Vancouver, 231 p. MILLER, D. J., and R. N. LEA. 1972. Guide to the coastal marine fishes of California. Calif. Dep. Fish Game, Fish. Bull. 157, 235 p. MISITANO, D. A. 1978. Description of laboratory-reared larvae of the buffalo sculpin, Enophrys bison Girard (Pisces:Cottidae). Copeia 1978:635- 642. MORRIS, R. W. 1951. Early development of the cottid fish, Clinocottus recalvus (Greeley). Calif. Fish Game 37:281-300. NELSON, J. S. 1977. Fishes of the southern hemisphere genus Neophrynichthys (Scorpaenifosmes : Cottoidei), with descriptions of two new spe- cies from New Zealand and Macquarie Island. J. R. Soc. N.Z. 7:485-511. O'CONNELL, C. P. 1953. The life history of the cabezon Scorpaenichthys marmoratus (Ayres). Calif. Dep. Fish Game, Fish. Bull. 93:1-76. OKIYAMA, M., and H. SANDO. 1976. Early life history of the sea raven, Hemitripterus uillosus, (Hemitripterinae, Cottidae) in the Japan Sea. [In Jpn., Engl, abstr.] Bull. Jpn. Sea Reg. Fish. Res. Lab 27:1-10. PEDEN, A. E. 1964. A systematic revision of the Hemilepidotinae, a subfamily of the cottid fishes. M.S. Thesis, Univ. British Columbia, Van- couver, 162 p. QUAST, J. C, and E. L. HALL. 1972. List of fishes of Alaska and adjacent waters with a guide to some of their literature. U.S. Dep. Commer., NOAA Tech. Rep. NMFS SSRF-658, 47 p. RASS, T. S. 1949. Fish eggs and larvae of the Barents Sea. Proc. All Union Res. Inst. Mar. Fish. Oceanogr. (VNIRO) 17:9-65. RICHARDSON, S. L. In press. Current knowledge of northeast Pacific sculpin larva (Family Cottidae) with notes on relationships within the family. ICES/Early Life History Symposium, Cons. Int. Expl. Mer., Rapp. Proc. -Verb. Reunions. RICHARDSON, S. L., and W. A. LAROCHE. 1979. Development and occurrence of larvae and juveniles of the rockfishes Sebastes crameri, Sebastes pinniger, and Sebastes helvomaculatus (Family Scorpaenidae) off Oregon. Fish. Bull., U.S. 77:1-46. 55 RICHARDSON. S. L., and W. G. PEARCY. 1977. Coastal and oceanic fish larvae in an area of upwelling off Ya- quina Bay, Oregon. Fish. Bull., U.S. 75:125-145. RUSSELL. F. S. 1976. The eggs and planktonic stages of British marine fishes. Acad. Press, N.Y., 524 p. SANDERCOCK, F. K., and N. J. WILIMOVSKY. 1968. Revision of the cottid genus Enophrys. Copeia 1968:832- 853. SCOTT, W. B., and E. J. CROSSMAN. 1973. Freshwater fishes of Canada. Bull. Fish. Res. Board Can., Bull. 184, 966 p. STEIN, D. L., and C. E. BOND. 1978. A new deep-sea fish from the eastern north Pacific, Psychro- lutes phrictus (Pisces: Cottidae [Psychrolutinae]). Los Ang. Cty. Mus., Contrib. Sci. 296, 9 p. STEIN, R. 1972. Identification of some Pacific cottids. M.S. Thesis, Hum- boldt State Coll., Areata, Calif., 41 p. 1973. Description of laboratory-reared larvae of Oligocottus macu- losus Girard (Pisces:Cottidae). Copeia 1973:373-377. TAYLOR, W. R. 1967. An enzyme method of clearing and staining small verte- brates. Proc. U.S. Natl. Mus. 122(3596):1-17. WARFEL, H. E., and D. MERRIMAN. 1944. The spawning habits, eggs and larvae of the sea raven, Hemi- tripterus americanus, in southern New England. Copeia 1944:197-205. WHITE, W. A. 1977. Taxonomic composition, abundance, distribution and seasonality of fish eggs and larvae in Newport Bay, Cali- fornia. M.S. Thesis, Calif. State Univ., Fullerton, 107 p. 56 NOAA TECHNICAL REPORTS NMFS CIRCULAR AND SPECIAL SCIENTIFIC REPORT— FISHERIES GUIDELINES FOR CONTRIBUTORS CONTENTS OF MANUSCRIPT First page. Give the title (as concise as possible) of the paper and the author's name, and fo ;note the author's affiliation, mailing address, and ZIP code. Contents. Contains the text headings and abbreviated figure legends and table headings. Dots should follow each entry and page numbers should be omitted. Abstract. Not to exceed one double-spaced page. Footnotes and literature citations do not belong in the abstract. Text. See also Form of the Manuscript below. Follow the U.S. Government Printing Office Style Manual, 1973 edition. Fish names, follow the American Fisheries Society Special Publica- tion No. 6, A List of Common and Scientific Names of Fishes ]rom the United States and Canada, third edition, 1970. Use short, brief, informative headings in place of "Materials and Methods." Text footnotes. Type on a separate sheet from the text. For unpublished or some processed material, give author, year, title of manuscript, number of pages, and where it is filed — agency and its location. Personal communications. Cite name in text and footnote. Cite in footnote: John J. Jones, Fishery Biologist, Scripps Insti- tution of Oceanography, La Jolla, CA 92037, pers. commun. 21 May 1977. Figures. Should be self-explanatory, not requiring reference to the text. All figures should be cited consecutively in the text and their placement indicated in the left-hand margin of the manuscript. Photographs and line drawings should be of "professional" qual'ty — clear and balanced, and can be re- duced to 6 'A inches (40 picas) for page width or to 3 l A inches (19 picas) for single-column width, but no more than 9 inches (54 picas) high. Photos should be printed on glossy paper — sharply focussed, good contrast. Label each figure. List, and typed dou- ble spaced, each figure legend. DO NOT SEND original figures to the Scientific Editor; NMFS Scientific Publications Office will request these if they are needed. Tables. Each table should start on a separate page and should be self-explanatory, not requiring reference to the text. Headings should be short but amply descriptive. Use only horizontal rules. Number table footnotes consecutively across the page from left to right in Arabic numerals; and to avoid con- fusion with powers, pit je them to the left of the numerals. If the original tbles are typed in our format and are clean and leg- ible, these tables will be reproduced as they are. In the text all tables should be cited consecutively and their placement indi- cated in the left-hand margin of the manuscript. Acknowledgments. Place at the end of text. Give credit only to those who gave exceptional contributions and not to those whose contributions are part of their normal duties. Literature cited. In text as: Smith and Jones (1977) or (Smith and Jones 1977); if more than one author, list according to years (e.g., Smith 1936; Jones et al. 1975; Doe 1977). All papers re- ferred to in the text should be listed alphabetically by the senior author's surname under the heading "Literature Cited"; only the author's surname and initials are required in the author line. The author is responsible for the accuracy of the literature cita- tions. Abbreviations of names of periodicals and serials should conform to Biological Abstracts List of Serials with Title Abbre- viations. Format, see recent SSRF or Circular. Abbreviations and symbols. Common ones, such as mm, m, g, ml, mg, °C (for Celsius), %, %„, etc., should be used. Abbrevi- ate units of measures only when used with numerals; periods are rarely used in these abbreviations. But periods are used in et al., vs., e.g., i.e., Wash. (WA is used only with ZIP code), etc. Abbreviations are acceptable in tables and figures where there is lack of space. Measurements. Should be given in metre units. Other equivalent units may be given in parentheses. FORM OF THE MANUSCRIPT Original of the manuscript should be typed double-spaced on white bond paper. Triple space above heading: Send good duplicated copies of manuscript rather than carbon copies. The sequence of the material should be: FIRST PAGE CONTENTS ABSTRACT TEXT LITERATURE CITED TEXT FOOTNOTES APPENDIX TABLES (each table should be numbered with an Arabic numeral and heading provided) LIST OF FIGURE LEGENDS (Entire figure legends, includ- ing "Figure" before each number) FIGURES ADDITIONAL INFORMATION Send ribbon copy and two du. Heated copies of the manuscript to: Dr. Jay C. Quast, Scientific Editor Northwest and Alaska Fisheries Center Auke Bay Laboratory National Marine Fisheries Service, NOAA P.O. Box 155 Auke Bay, AK 99821 Copies. Fifty copies will be supplied to the senior author and 100 to his organization free of charge. UNITED STATES DEPARTMENT OF COMMERCE •MTIONAl OCEANIC AND ATMOSPHERIC ADMINISTRATION NATIONAL MARINE FISHERIES SERVICE SCIENTIFIC PUBLICATIONS STAFF ROOM 450 1107 N E 45TH ST SEATTLE, WA 98105 PENN STATE UNIVERSITY LIBRARIES OFFICIAL BUSINESS ADDDD72DnD7fl BULK RATE NOAA SCIENTIFIC AND TECHNICAL PUBLICATIONS NOAA, the National Oceanic and Atmospheric Administration, was established as part of the Department of Commerce on October 3, 1970. The mission responsibilities of NOAA are to monitor and predict the state of the solid Earth, the oceans and their living resources, the atmosphere, and the space environment of the Earth, and to assess the socioeconomic impact of natural and technological changes in the environment. The six Major Line Components of NOAA regularly produce various types of scientific and technical infor- mation in the following kinds of publications: PROFESSIONAL PAPERS— Important definitive research results, major techniques, and special in- vestigations. TECHNICAL REPORTS— Journal quality with extensive details, mathematical developments, or data listings. TECHNICAL MEMORANDUMS— Reports of preliminary, partial, or negative research or tech- nology results, interim instructions, and the like. CONTRACT AND GRANT REPORTS— Reports prepared by contractors or grantees under NOAA sponsorship. TECHNICAL SERVICE PUBLICATIONS— These are publications containing data, observations, instructions, etc. A partial listing: Data serials; Pre- diction and outlook periodicals; Technical manuals, training papers, planning reports, and information serials; and Miscellaneous technical publications. 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