sign in sign up
<<
Home , Artedius, Artedius harringtoni, Blepsias, Blepsias cirrhosus, Clinocottus, Clinocottus acuticeps

CURRENT KNOWLEDGE OF LARVAE OF (PISCES: AND ALLIES) IN NORTHEAST PACIFIC GENERA

WITH NOTES ON INTERGENERIC RELATIONSHIPS 1

SALLY L. RICHARDSON2

ABSTRACf

Current knowledge of cottid larvae in northeast Pacific genera is summarized. Larvae are known for representatives of25 ofthe 40 genera reported from Baja California to the Aleutian Islands although two genera, and lcelus, are represented only by which live in other areas as adults. Included are illustrationsoflarvaeof29species representing the 25 genera plus one potentially new northeast Pacific , identified only as "Cottoid Type A." The larvae exhibit a wide diversity of form. Based on shared larval characters, including spine patterns, body shape, and pigmentation, 6 phenetically derived groups of genera are apparent within the 25 genera for which representatives are considered: 1) A rtedius, , , Orthonopias; 2) Paricelinus, , lcelus, Chitonotus, ; 3) Dasycottus, Psychrolutes, Gilbertidia, ?Malacocottus, "Cottoid Type A"; 4) Scorpaenichthys, ; 5) , ; 6) Leptocottus, . Six genera do not fit with any group: , Gymnocanthus, , , Rhamphocottus, . Ifthese preliminary larval groupings reflect relationship, as evidence indicates, they tend to support a number of previously implied relationships within the cottids, but there are some important differences. These include the distinctiveness of the (Group 1) line; the separation of Artedius and Icelus, once considered closely related; the relationship of Paricelinus, generally considered a primitive and rather distinct form, with other members of Group 2; the apparent relationship oflcelus to other genera in Group 2 and its questionable placement in a separate family; the distinctiveness of Radulinus, previously considered to be related to Chitonotus and Icelirms.

The Cottidae, which in this paper are considered Aleutian Islands (Table 1). With the recent work broadly to include sculpinlike of Cottidae, by Richardson and Washington (1980), larvae are Icelidae, , Comephoridae, Nor­ now known for representatives of 25 of these 40 lllanichthyidae, Cottunculidae, and Psychrolut­ cottid genera, although two genera, Icelus and idae ofthe suborder Cottoidei ofGreenwood et al. Gymnocanthus, are represented only by larvae of (1966), comprises a diverse group oftemperate and species that live in other areas as adults. boreal fishes. Nelson (1976) estimated that the The purpose ofthis paper is twofold. It presents group may contain over 350 species, three-fourths for the first time a summary of important cottid lllarine, in about 86 genera. They are generally larval characters (those characters occurring only Coastal fishes inhabitingall oceans but the Indian. during the larval period and most useful in identi­ Greatest species diversity occurs in the North fying and distinguishing species) based on the Pacific. The systematics of the group are not well larvae of these 25 northeast Pacific genera. (Lar­ understood (Quast 1965; Nelson 1976). vae of these genera that are known for species Until recently, larvae of relatively few cottids inhabiting other areas as adults are also con­ ?ad been described. They were a difficult group to sidered.) This knowledge, which is a necessary Identify in ichthyoplankton collections, particu­ prerequisitefor systematic studies usinglarvae, is larly in the northeast Pacific where 40 genera are presented to provide a foundation to which future reported to occur between Baja California and the work on cottid larvae can be compared and upon --;:----- which it can be expanded as more larvae become ''!-'his paper was presented at the Second International Sym- known. The paper also presents a preliminary f1~um on The Early Life History of (sponsored by ICES, 197 ,ICNAF, IAOB, SCOR) held at Woods Hole, Mass., 2-5 April examination of generic groupings within these 91: An abstract of the paper appeared in the symposium northeast Pacific cottid genera based on shared P~b 'cation. S

TABLE I.-List ofcottid genera occuring in the northeast Pacific Ocean between Baja California and the Aleutian Islands based on Howe and Richardson (text footnote 3) with a summary of illustrations (accessible to author) oflarvae known for those genera world­ wide. Lengths (millimeters) oflarvae are reported as they appeared in the literature: NL = notochord length; 8L = standard length; TL = total length; mm = no length definition was given. Genus and species Reference Sizes illustrated Alfediel/us ­ Altedius harringtoni Richardson and Washington 1980 3.0,4.7,6.9 mm NL, 7,3, 9.3,13.6 mm SL A. laferalis Budd 1940 4,1 mm SL Marliave 1975 4,8,11,14 mm TL Alfedius Type 2 Richardson and Washington 1980 3.0,4,7,6,0 mm NL, 7,2, 9.9,11.8 mm SL Ascel/chthys ­ Asemichthys ­ Biepsias ci"hosus Blackburn 1973 12.2mm SL Marliave 1975 10,14,19,15.5 mm TL Chitono/us pugetensis Richardson and Washington 1980 3.0,6.3 mm NL, 8.5, 11.5, 15.4, 16.6 mm SL Misitano 1980 3.0, 4.8 mm SL Richardson and Washington 1980 3.7,3.9,6.9 mm NL, 7.6,10.4,13.8,16.5 mm SL C. analis Eigenmann 1892 ca.4mm Budd 1940 ca.4mm C. reca/vus Morris 1951 4.6,5.0,7.6,8.3,9.9,10.8,18.0,24.3 mm TL Cottus asper Stein 1972 5.5,9.0,10.8 mm TL Richardson and Washington 1980 5.2 mm NL, 8.2, 9.9 mm SL Dasycottus setiger Blackburn 1973 7.4 mm SL Blackburn 1973 7.5mmSL Marliave 1975 10 mm TL Misitano 1978 5.0,5.4, 5.8, 6,7, 7.1,7.6 mm SL Richardson and Washington 1980 4.8, 7.0 mm NL, 9.1 mm SL E. bubalis' Cunningham 1891" 5.7mm Mcintosh and Masterman 1897" Larva (size not given) Ehrenbaum 1904" 5.8,10,11 mm Ehrenbaum 1905-9 5.8,10,11 mm Russell 1976' 4.5,5.7,6.4,9.5 mm E. liI/jeborgi' Bruun 1925' 5.6,6.8,8.7 mm TL Rass 1949' 6.8mm Russell 19766 4.08,4.2,5.7,7.0 mm Eurymen- GI/bart/dia sigalutes Blackburn 1973 7.9,9.5 mm SL Marliave 1975 7,13,15,25,34 mm TL Gymnocanthus herzenstainl' Kyushln 1970 5.79,6.59,7.55 mm G. tricuspis 1 Koefoed 1907 10.7,12.7,15.5 mm Rass 1949 9mm Khan 1972 12.2,13.9,15.9 mm TL G. ventralis 1 Ehranbaum 1905-9 15,18 mm Heml/epidotus gilberti' Gorbunova 1964 7.5,11.4,17.5 mm Hattori 1964 7.1,11.6,19.2,24.8,32.5 mm H. heml/epidotus Gorbunova 1964 7.25,10.5 mm Peden 1978 ca.20mm SL Richardson and Washington 1980 5.8,5.9,9.1 mm NL, 10.7, 11.5, 19.0 mm SL H.jordani Gorbunova 1964 6.4,10.7,13.0 mm Peden 1978 ca. 20 mm SL H.papilio Gorbunova 1964 10.7,13.7mm H. spinosus Follett 1952 12,21 mm SL Peden 1978 ca. 20 mm SL Richardson and Washington 1980 5.0, 6.6, 8.9 mm NL, 11.0; "11.8, 19.0 mm SL H. zapus Pedan 1978 ca.20mmSL ' Warfel and Merriman 1944 ca,12mm Khan 1972 11.7,' 14.5, 18.8 mm TL Fuiman 1976 12.6,15,5,20 mm TL H. viliosus· Kyushln 1968 14.78,15.57,16,52 mm SL Okiyama and sando 1976 11.6,14.4,17.4,20.0 'llm leal/nus spp.· Richardson and Washington 1980 3.3,8.6 mm NL, 10.9, 13.8, 15.2, 16.5, 12.5,16.6 mm SL tee/us bicornis' Ehrenbaum 1905-9'· 25mm Rass 1949 12.3mm ­ Laiocottus ­ Leptocottus armatus Jones 1962 ca. 4 mm Blackburn 1973 7,6,8.3,12.0 mm SL Marllave 1975 8,12,13mmTL WMe1977 4.9mmNL Richardson and Washington 1980 5.1,8.1 mm NL, 11,1 mm SL Ma/acocottus 1M. zonurus Type 1 Richardson and Bond 11 7.0,9.8, 14.2,24.0 mm SL Richardson unpubl. data 6.6,7.0,8.8,9.8,10.4,14.2,24.0 mm SL ' Perlmutter 1939 6mm Khan 1972 5.0,7.1,9.7, 11.8 mm TL Lund and Marcy 1975 5.4,6,1,6,8,7.5,8.5,9.2 mm TL M. octodecemsplnosus' Colton and Marak 1969 6.8,8.5,10.5,15.2 mm TL Khan 1972 7,0,9,5,10.7, 12.5, 14.5 mm TL M. polyacanfhocephalus Blackburn 1973 7.7,10.7 mm SL M. quadricornis 1 (marine form) Zvjagina 1963 12.3, 12.8, 13.6, 14.5, 16.2,32 mm Khan 1972 12.8,14.5,17.0 mm TL Khan and Faber 1974 12,8,14.4,17.0 mm TL 104 RICHARDSON: CURRENT KNOWLEDGE OF LARVAE

TABLE I.-Continued.

Genus and species Reference Sizes illustrated M. scorpius' Mcintosh and Prince 1890" Larva (size not given) Mcintosh and Masterman 1897" Larva (size not given) Ehrenbaum 1904" 8.24,8.16,10,18 mm Ehrenbaum 1905-9 8.2,10,18 mm Koefoed 1907 9.5mm Rass 1949 7.9,9.3 mm Bigelow and Schroeder 1953 8.2,10,18mm Khan 1972 7.6,8.5,10.4,14.0,17.4 mm TL Russell 1976 7.5,9.5,10,14 mm Nautichthys oculo/asciatus Blackburn 1973 7.5,13 mm SL Marliave 1975 9.5,13,17,26 mm TL Richardson and Washington 1980 11.7 mm NL, 16.6 mm SL Otigocottus maculosus Stein 1972 4.6-5.2,6.0,6.6, 9.2 mm TL Stein 1973 4.6-5.2,6.0,6.6,9.2 mm TL O. snyderi Stein 1972 4.5-4.75,5.5 mm TL Orthonoplas triacis Bolin 1941 ca. 3-4 mmSL Par/cetinus hoptiticus Richardson and Washington 1980 5.6,6.2 mm NL, 13.8, 18.6,25.6 mm SL Phal/ocottus - - Psychrolutes paradoxus Blackburn 1973 10.3mm SL Marliave 1975 10.5,13,14,13 mm TL Radul/nus asprel/us Richardson and Washington 1980 4.7,7.9,9.6,10.9 mm NL, 12.6, 14.4 mm SL R. boleo/das Richardson and Washington 1980 8.7 mm NL Rhamphocottus richardsonl Blackburn 1973 6.7,10 mm SL Marliave 1975 10,11.5,15 mm TL Richardson and Washington 1980 8.4 mm NL, 10.6, 11.7 mm SL Scorpaenichthys marmoratus O'Connell 1953 5.85,6.26, 10, 17, 30, 48 mm Richardson and Washington 1980 5.3,7.5,8.6 mm NL, 8.7,10.4,13.8 mm SL S/gm/stes- Ste/gistrum - Sternias - SUag/cottus - Synch/rus _ Thyr/s/cus _ Trlglops murray/' Khan 1972 8.4, 11.6, 18.9, 23.4 mm TL T. p/ngall' Ehrenbaum 1905-9 18mm Koefoed 1907 13,16.5,22 mm Rass 1949 10mm Tr/g/ops sp. Blackburn 1973 8.3, 12mm SL Richardson and Washington 1980 6.9 mm NL, 15.4 mm SL Zastlce/us _

-;Not northaast Pacific species. 'Probably (Laroche textlootnote 6) . •As Cottus bubel/so 'Species occurs in northeast Pacific but larvae described from other areas. ,As Tauru/us bubelis. 'Not identified below genus level. ,As Cottus /II/jaborg/. '0As Centriderm/chthys hamatus . ,As Acanthocottus /II/jeborgi. "Textlootnote 5. As Taurulus /ll/jeborg/. "As Cottus scorpius. in reducing taxonomic problems. The potential pigmentation. Although meristic characters may usefulness of the larval groups in providing in­ be of prime utility in identifying cottid larvae, sights into systematic relationships and evolu­ they persist in adults and are not considered truly ~ionary trends within this difficult group offishes larval. Meristic characters for northeast Pacific IS also discussed. The use of larval forms of cottids have been discussed by Richardson and fishes to elucidate systematic relationships has Washington (1980) and Howe and Richardson.3 been demonstrated in a number of groups, e.g., The purpose of this summary is to point out the ceratioids (Bertlesen 1941), myctophids (Moser kinds of larval characters that are useful for and Ahlstrom 1970, 1972, 1974), gonostomatids identification and the spectrum in which those (Ahlstrom 1974), scombroids (Okiyama and characters may be exhibited since cottid larvae DeYanagi 1978), and serranids (Kendall 1979). manifest a wide diversity ofform. This summary is based on only the representa­ LARVAL CHARACTERS OF COTTIDS tives of northeast Pacific cottid genera listed in

This is a summary of important larval charac­ "Howe, K. D., and S. L. Richardson. 1978. Taxonomic review and meristic variation in marine sculpins (Osteichthys: ters in cottids, i.e., those characters occurringonly Cottidae) of the northeast Pacific Ocean. Final rep" NOAA ?uring the larval period which can be of most use NMFS Contract No. 03-78-M02-120, 1 January 1978 to 30 September 1978, 142 p. Northwest and Alaska Fisheries Center, In distinguishing species. These characters in­ National Marine Fisheries Service, NOAA, 2725 Montlake clude preopercular spine pattern, body shape, and Boulevard East, Seattle, WA 98112. 105 FISHERY BULLETIN: VOL. 79, NO.1 Table 1. Species from areas outside the northeast rounded (Scorpaenichthys, Hemilepidotus) or Pacific are included when larvae are known be­ pointed (lcelinus, Chitonotus). Snout to anus cause of the taxonomic information their larvae length can be rather short, < 40% SL (standard may provide. Generic level designations are used length) (Dasycottus), to moderately long, >60% throughout the text for continuity and emphasis SL (Rhamphocottus), although this can change although larvae of all species (number of species with development. The gut may appear tightly based on the taxonomic status summary by Howe compacted (Dasycottus) or be distinctively coiled and Richardson footnote 3) in a genus may not be (Cottus). The hindgut may trail somewhat below known. In some cases the genera are monotypic the body (Artedius, Clinocottus). Unusual gut (Chitonotus, Dasycottus, Gilbertidia, Leptocottus, diverticula may be present (Artedius, Clinocot­ Orthonopias, Paricelinus, Rhamphocottus, Scor­ tus). Pectoral fins may be noticeably elongated paenichthys) and thus larval characters of the (Nautichthys) or fanlike early in development genus may readily be defined. At least some (Myoxocephalus). developmental stages are known for all six species Melanistic pigment patterns range from rela­ of Hemilepidotus, providing good generic level tively unpigmented to heavily pigmented. Pig­ definition. Insome cases larvaeofa few, but not all ment may be variously present or absent over the species within a genus are known [Artedius (3 head, snout, cheek, jaws, cleithral base, throat. species out of7); Gymnocanthus (3 of6); Hemitrip­ Pigment over the dorsolateral surface of the gut terus (2 of 2 or 3); Myoxocephalus (5 of 18); may vary in intensity, ventrolateral extent, and Oligocottus (2 of4); Radulinus (2 of5); Triglops (3 pattern (e.g., bars, Leptocottus; distinct round of 9)]. In those instances, constancy of larval melanophores, Enophrys). In some species the characters among species provides good indica­ entire gut region is pigmented (Paricelinus). The tions ofgeneric level definition. Larvae ofIcelinus ventral midline ofthe gut may have a distinct line spp. have only been described at the generic level ofmelanophores (Cottus, some Myoxocephalus) or as none of the eight species have yet been distin­ be unpigmented (Scorpaenichthys). The nape may guished. For some genera, larvae are known for be distinctively pigmented (Artedius, Enophrys). only one of a few species: Blepsias (l of 2), Cottus The lateral body surface above the gut may be (1 of 2 brackish water species), Icelus (l of 13), unpigmented (Chitonotus), have dorsolateral pig­ Malacocottus (1 of 5), Nautichthys (1 of 3), ment not extending to the gut (Radulinus) or be Psychrolutes (1 of2). In those, generic level defini­ entirely pigmented (Scorpaenichthys). In the tail tion may not be as precise; however, larvae of all region posterior to the anus, pigment may be species appear rather distinctive and thus may be absent (some Triglops, Dasycottus), present along good representatives oftheir genera. Inthe follow­ only the ventral midline (Artedius, Chitonotus), ing summary those genera which provide the best present along only the ventral and dorsal midlines examples of patterns are listed in parentheses. [Gymnocanthus, small (< 8 mm) Hemilepidotus], Principal preopercular spines typically (18 of25 or present on the lateral body surface, sometimes genera) number 4 (Scorpaenichthys, Icelinus, in combination with a ventral midline series Leptocottus, Enophrys) and may vary in degree of (Scorpaenichthys, Radulinus, Blepsias). Num­ development. Modifications of this basic pattern ber, spacing, position, and shape of ventral mid­ may occur (Myoxocephalus, ?Malacocottus) in line melanophores are important as is the pos­ which four main spines are present with one or terior extent of lateral pigment. Melanophores two auxiliary spines. Another pattern consists of may variously appear along the caudal fin base multiple preopercular spines, usually small, num­ (Paricelinus, Chitonotus). Pectoral fins are gen­ bering up to ca. 25 (Artedius, Clinocottus). Some­ erally unpigmented, but some species have heav­ times only one spine is present (Rhamphocottus) ily pigmented fins (Psychrolutes, Gilbertidia) or a or none (Psychrolutes, Gilbertidia). Spines in pigment band along the fin margin (Nautichthys). other regions of the head (particularly parietal­ nuchal, postocular, posttemporal-supracleithral, LARVAL COTTID GROUPS opercular) may also be important. General body shape can range from rather Within the 25 cottid generaconsidered, 6 groups stubby and deep (Artedius, Enophrys) to moder­ of genera are apparent based on shared larval ately slender and elongate (lcelinus, Triglops) to characters, Le., similarity, and 6 genera do not fit globose (?Malacocottus). The snout can be quite into any group (Table 2). Characters within each 106 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE

TABLE 2.-Groupings of 25 cottid genera reported to occur in the northeast Pacific Ocean between Baja California and the Aleutian Islands based on shared larval characters. Group characteristics were based on representative species for which larvae are known, as listed in Table 1. Also included in the groupings is an unidentified larval type, "Cottoid Type P;' of Richardson and Washington (1980) ~hich may represent a new genus. ~oup General characteristics Genera Multiple preopercular spines, rounded snout, stubby shape, slightly trailing gut, Artedius, Clinocottus, Oligocottus, sometimes with gut protrusions or diverticula Orthonoplas 2 Four preopercular spines, pointed snout, moderately slender, postanal pigment when Paricel/nus, Trlglops, Icelus, present usually restricted to ventral midline Chitonotus, leel/nus 3 Four principal preopercular spines or none, rounded snout, often globose shape Dasyeottus, Psyehrolutes, Gllbertldia, with loose skin, pigmented pectoral fins ?Malaeoeottus, Cottoid Type A (new genus?) 4 Four preopercular spines, rounded snout, relatively deep bodied, ca. 4-5 mm NL at Seorpaenlehthys, Hemi/epidotus hatching, postanal pigment dorsally, ventrally, and laterally 5 Four preopercular spines not pronounced, rounded snout, relatively slender, post· Blepsias, Nautiehthys anal pigment dorsally, ventrally, laterally, probably >7 mm NL at hatching, pectoral fins unpigmented or with pigment band near margin 6 Four preopercular spines, rounded snout, relalively slender, no additional head Leptoeottus, Cot/us spines, postanal pigment restricted to ventral midline ~roUPed genera Enophrys, Gymnocanthus, Myoxocephalus, Radulinus, Rhamphocottus, Hemitripterus group and of each ungrouped genus are summa­ ber of spines (e.g., none, two, cluster) in the rized to facilitate recognition and minimize tax­ parietal and posttemporal-supracleithral regions; onomic and identification problems involving presence or absence of pigment on the nape or cottid larvae. These groupings are based on com­ head. plete developmental series to theextent available, Although the multiple preopercular spine pat­ but only representative figures illustrating one tern persists through the larval period, adults point on a developmental continuum are pre­ have four preopercular spines with the lower three ~ented (Figures 1-9). The groupings are necessar­ reduced and the upper variously modified. Rem­ lly preliminary because not all species in all nants of the larval serrations have been observed 4 genera are known as larvae. The groups described only in adult A. notospilotus (Howe ). It is unclear below are not arranged in any particular order. which four spines of the larvae persist in adults. Generic designations are used as discussed in the previous section, Group 2

Group 1 This is also a rather cohesive group (Figure 3) consisting of slender forms with pointed snouts This is the tightest group among the 25 genera. and four prominent preopercular spines [Paricel­ InclUded are Artedius, Clinocottus, Oligocottus, inus, Triglops, [celus (tentatively), Chitonotus, and tentatively Orthonopias (Figures 1, 2). The Icelinus]. This general body shape is remarkably Unique multiple preopercular spine pattern dis­ similar among genera and is not found in any tinguishes it from all other groups or genera. other genera considered. All have a relatively [Although a complete series of Orthonopias has short snout to anus distance. Postanal ventral not been described and the spine pattern is un­ midline pigment is usually present (absent in one known, small larvae (Figure 2) are very similar species of Triglops) with some additional melano­ to Artedius in form and pigment characteristics phores along the caudal fin base. Dorsal midline and are tentatively included in this group.] The pigment is usually absent except for a few spots in stUbby body shape, rounded snout, and somewhat some Icelinus and possibly a row in some late trailing gut are remarkably consistent within the stage Triglops. Generic differences include degree group. Presently, identification to genus based on of gut pigmentation (e.g., darkest in Paricelinus larval characters is still difficult and in need of and some Triglops), number and position of ven­ better definition. Characters used to distinguish tral midline melanophores, and degree of head species (besides fin ray counts) include: number, spination (e.g., postocular spines in Paricelinus Spacing, and shape of ventral midline melano­ and Triglops). Phores; intensity ofgut pigmentation; presence of 4K, D, Howe, Ph.D. candidate, Department of Fisheries and unUsual gut diverticula; total number ofpreoper­ Wildlife, Oregon State University, Corvallis, OR 97331, pers. cUlar spines and position of largest spines; num- commun. September 1978. 107 FISHERY BULLETIN: VOL. 79. NO.1

c

FIGURE I.-Larvae ofA) (7.3 mm SL), B) Artedius Type 2 (9.9 mm SL), C) Clinocottus acuticeps (7.6 mm SL). D) Oligocottus maculosus (9.2 mm TL) (A-C, Richardson and Washington 1980; D, Stein 1973). 108 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE A

;.\\\~\ .... "",\ '\~.\\'.. "'\.\

--- ~- .' -_. -.. ~ -. B ~~":--~";;'-:':.'.'-~.'"

(;>..<_..J~~~~~--""--- . .

FIGURE 2.-Larvae of A) Artedius harringtani (3.0 mm NL) and B) Orthanapias triacis (""4 mm) (A, Richardson and Washington 1980; B, Bolin 1941).

The tentative placement of [celus with this often considered a separate family (Nelson 1976). group is of interest as it has been considered to ["Cottoid Type N' may possibly be Psychrolutes Constitute a distinct family, the Icelidae (Jordan phrictus but positive identification awaits addi­ 1923; Greenwood et al. 1966) based on the presence tional specimens-see Discussion by Richardson of scales in adults. Although larvae of [celus are and Washington (1980). If it is P phrictus, larval known to the author only from descriptions in the evidence indicates that the species is incorrectly literature (Table 1), they strongly resemble other placed and that a new northeast Pacific genus of Illembers of this group in form and preopercular cottid is in need of description.] This group is not spine pattern. Inclusion of Paricelinus is also of as cohesive as the two previous groups. The most interest as it has been considered to be a rather distinctive character of Group 3 is the pattern of distinct and primitive form (Bolin 1947). It pos­ pigmentation of the pectoral fin, a pattern not sesses five pelvic soft rays, the ancestral condition, found in any ofthe other genera considered. In all, Whereas the number ofsoft rays is reduced tothree at least the basal portion of the fin develops or two in other members ofthe group. pigment with the entire fin pigmented in Psychro­ lutes, Gilbertidia, and small « 9 mm SL) ?Mala­ Group 3 cocottus. Pigment on small (< 8 mm SL) Dasycot­ tus is restricted to the inside surface ofthe pectoral Group 3 (Figure 4) consists ofthe "psychrolutid" fin base but later it develops distally on the outer cottids [Dasycottus, Psychrolutes, Gilbertidia, surface. (The pigment band near the margin ofthe ?lvtalacocottus,S "Cottoid Type N' (new genus?)] elongated pectoral fin of Nautichthys is a very different pattern.) Only Dasycottus and ?Mala­ ~entification of larvae is tentative pending resolution of cocottus develop four preopercular spines, the :xonomic problems of adults at the generic level [see Howe and Blchardson footnote 3 and also Richardson, S. L., and C. E. latter genus with an accessory spine at the base of Pon.d. 1978. 'I\vo unusual cottoid fishes from the northeast the second spine. All but the more slender Dasy­ aciflc. Unpubl. manuscr., 6 p. + 25 figs. (Available from senior have relatively rounded snouts and deep ~uthor.) (Paper presented at the American Society of Ichthyolo­ cottus sts and Herpetologists, 1978.» bodies. Both?Malacocottus and "Cottoid Type N' 109 FISHERY BULLETIN: VOL. 79, NO. I

B

FIGURE 3.-LarvaeofA) (13.8 mm SL), B) Triglops sp. (15.4 mm SL), C) Icelus bicomis (25 mm), D) ChUanatus pugetensis (11.5 mm SL), E) Icelinus sp. (13.8 mm SL) (A, B, D, E, Richardson and Washington 1980; C, Ehrenbaum 1905·9).

110 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE ------~~r-)

;' / ~..:::.3Y~'--- ~-J~~

c .:/~

• Ii' ~ .~ .....

FIGURE 4.-Larvae of A) Dasycottus setiger (8 mm NL), B) Psychrolutes paradoxus (transfonning, 13 mm SL), C) Gilbertidia ~ga~utes 03 mm SL), D) ?Malacocottus ZOIlUruS 00.4 mm SLl, E) Cottoid Type A (9.8 mm SL) (A-D, original illustrations; ,Richardson and Washington 1980).

111 FISHERY BULLETIN: VOL. 79, NO. I develop a pronounced globose appearance with an snouts, and relatively deep bodies with rather outer bubble of skin. Small larvae of all forms heavy pigmentation, except at the smallest sizes have head and gut pigment and "Cottoid Type A" (Scorpaenichthys, Hemilepidotus) , Differences also has lateral pigment posterior to the anus. between genera include the longer gut and the Some lateral pigment develops later in all but preanal fin fold of Scorpaenichthys and the in­ Dasycottus. "Cottoid Type N' has unique "thumb­ creased head spination ofHemilepidotus (parietal, tack" prickles covering the belly region. The nuchal, postocular, posttemporal-supracleithral). pelvic fins of ?Malacocottus and "Cottoid Type N' Pigmentation is generally heavier in Scorpae­ often appear to be withdrawn into pockets ofskin. nichthys than in Hemilepidotus of comparable size. It is initially concentrated along the dorsal Group 4 and ventral midlines, particularly in Hemilepi­ dotus, filling in laterally with development. Lar­ Group 4 (Figure 5) includes larvae with vae of both are neustonic. four preopercular spines, conspicuously rounded The two genera in this group are certainly more

FIGURE 5.-Larvae of A) Scorpaenichthys m.armoratus (8.7 mm 3L), B) Hemilepidotus spinosus (11.0 mm 3L), C) H. hemilepidotus 00.7 mm 3L) (A-C, Richardson and Washington 1980). 112 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE similar to each other than to any other cottids ventral midline pigment along gut, and postanal considered, although Scorpaenichthys was given pigment restricted to ventral midline. Both hatch familial status in the past (Jordan 1923; Tarenets and transform at similar sizes, ca. 3-4 mm NL and 1941). ca. 10-12 mm SL, respectively. Leptocottus has a unique gut pigment pattern of bars, and Cottus Group 5 has a distinctively coiled gut.

This group consists of two elongated, slender­ Ungrouped Genera bodied genera, Blepsias and Nautichthys (Figure 6). Both hatch at a relatively large size, >7 mm Enophrys (Figure 8) has four pronounced pre­ NL (notochord length). Both have rounded snouts, opercular spines, rounded snout, deep stubby relatively heavy pigmentation, and four preoper­ shape, pigmented nape, and postanal pigment cular spines that never become pronounced and only along the ventral midline. It has a postocular sharp. Nautichthys (at least N. oculofasciatus) spine and opercular spines, and a preanal fin fold. develops greatly elongated pectoral fins soon after Melanophores over the gut are distinctively round hatching, each of which develops a pigment band in shape and densely concentrated. This suite of near its distal margin. larval characters is not shared by any other genus. The genera Blepsias and Nautichthys have Enophrys bears some resemblance to Group 6 been placed in a separate family, Blepsiidae, in (Leptocottus-Cottus) but differs too much to be the past (Jordan 1923). part of it. The deep body, bulging gut, and pig­ mented nape somewhat resemble Group 1 (Ar­ Group 6 tedius et a1.) but spine patterns differ drastically. Larvae of E. bubalis and E. lilljeborgi from the Group 6 (Figure 7) contains Leptocottus and North Atlantic (Table 1) are extremely similar to Cottus (as based on the brackish water species C. E. bison from the northeast Pacific. asper). They share several characters, including Gymnocanthus (Figure 8) apparently never de­ the relatively slender body, rounded snout, four velops pronounced preopercular spines, according Preopercular spines, absence ofother head spines, to the literature (Table 1). Larvae of G. tricuspis A

FIGURE G.-Larvae of A) Blepsias cirrhoslts (11 nun NL) and B) Nalttichthys ocltlofasciatlts (11.7 mm NL) (A, original illustration; B, Richardson and Washington 1980). 113 FISHERY BULLETIN: VOL. 79, NO. I

A .! " .... " .,' .' .. :......

.> .~' ..," ". ". ,. .~. ~ .'. '; ." .': :... .:' ~; .. ~.~.( ..~:..•:.;.:::~ :'~~"'::' •.~.-~

------...... 4,. ~ ...

B ,,'" " . " .: .: .... '

~-_..,..--J-

FIGURE 7.-Larvaeof A) Leplocoltlls annalus (8.1 mm NL) and B) Coitus asped8.2 mm SL) (A, B, Richardson and Washington 1980). develop three very small preopercular spines, fold is present (in some or maybe all species). The visible only on stained specimens, and the spines pattern of preopercular spination is unusual. All of 14-15 mm G. ventralis are barely perceptible. species apparently have four principal spines but Gymnocanthus is relatively slender with a semi­ some may have one or two posteriorly directed pointed snout; it develops dorsal (at least G. auxiliary spines anterior to and near the base of tricuspis), ventral, and lateral midline pigment. the second principal spine and/or an auxiliary Although it shares some characters with Myoxo­ spine ventral and adjacent to the fourth principal 6 cephalus, grouping the two does not seem war­ spine (Laroche ). Development of this unusual ranted. The spacing of the ventral series of pattern has not been adequately described for any midline melanophores somewhat resembles the species of Myoxocephalus and needs closer exam­ situation in Myoxocephalus. General shape of ination. Auxiliary preopercular spines are un­ species of the two genera is also similar although known in other genera except ?Malacocottus, Myoxocephalus is much spinier. Some figures which has an anteriorly directed accessory spine indicate that Gymnocanthus has a preanal fin fold at the base of the second preopercular spine. as do at least some Myoxocephalus. Radulinus (Figure 9) is a heavily pigmented Myoxocephalus (Figure 8) may have four to six form with four preopercular spines (never pro­ preopercular spines, two of which may be auxil­ nounced), relatively pointed snout, slender body iary. Larvae are moderately slender with a semi­ (R. boleoides is deeper bodied than R. asprellus), pointed snout. Two distinct larval forms (subgen­ and long gut. It shares the characters of pointed era?) occur in the genus. Those with dense lateral snout and slender body with Group 2 (ParicelinuS pigment in the region posterior to the anus (M. et al.) but differs too much (longer gut, heavier polyacanthocephalus, M. scorpius) and those with pigmentation, reduced prominence of preoper­ only ventral midline pigment postanally (M. cular spines) to be part of that group. It shares a aeneus, M. octodecemspinosus). Another relative­ few similarities with the dark species of Myoxo­ ly unpigmented species, M. quadricornis (marine cephalus, including a series of ventral midline form), develops a lateral midline pigment series. melanophores on the tail beyond the lateral pig­ The spacing of the ventral midline melanophores ment (in larvae <8-9 mm NL) and a relatively is similar in all species as is gut pigment. The fan­ GJ. L. Laroche, Research Assistant, Gulf Coast Research shaped pectoral fin is obvious early in develop­ Laboratory, East Beach Drive, Ocean Springs, MS 39564, pel's. ment (at least in some species) and a preanal fin cornmun. September 1978. 114 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE

A

". ' ..

~~\ ~~ o ..~\_--._)/.. / . ;;.' ;y/. . ..

. I - , + ... . III 7 .. , ... -.",\,~~ -~---~

~~G lIRE 8. _ Larvae of A) Enophrys bison (7.0 mm NL) l B) Gymnocanthus tricuspis (13.9 mm), C) Myoxocephalus aeneas (8.5 mm TL) I D !vI. polyacanthocephalus (12 nun SL) (A, Richardson and Washington 1980; B, Khan 1972; C, Lund and Marcy 1975; , Original illustration).

115 FISHERY BULLETIN: VOL. 79. NO. I A

FIGURE 9.-Larvae of A) Radulinus asprellus (10.9 nun NLl. B) Rhamphocottus richardsoni (8.4 nun NL). C) Hemitripterus uillosus (17.4 mm SL) (A, B. Richardson and Washington 1980; C. Okiyama and Sando 1976).

unpigmented area on the body above the abdom­ shape, but it has a longer gut, more pigment inal cavity. Small larvae also resemble Scar­ ventrally along the head and gut, and a pigmented paenichthys except that they have a distinct preanal fin fold. By 8-9 rom SL, the distinct shape lateral midline series of melanophores and soon of R hamphocottus is obvious and spinelike develop a pointed snout and more slender body. prickles develop over the body. The single species Rhamphacattus (Figure 9) is one of the most has been considered to represent a separate fam­ aberrant cottid forms. It develops only one pre­ ily (Jordan and Evermann 1898; Jordan 1923; opercular spine, an unusual snout, a deep body, Taranets 1941). heavy pigmentation, and a preanal fin fold. At Hemitripterus (Figure 9) is also a heavily pig­ small sizes, ca. 6-7 rom NL, it bears some resem­ mented and distinct form. Based on the literature blance to Scorpaenichthys in pigmentation and (Table 1), it has four preopercular spines, a moder- 116 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE ately pointed snout, relatively deep body, and long spine patterns, relative body form, and pigmenta­ gut. Larvae are quite large, ca. 12-14 mm NL, at tion have been used to demonstrate or clarify hatching. The larvae of H. americanus from the systematic relationships in other groups offishes, Atlantic and H. villosus from Japan (Table 1) are e.g., scorpelarchids (Johnson 1974), gonostomatids Very similar. The heavy body pigment is a char­ (Ahlstrom 1974), myctophids (Moser and Ahl­ acter shared with a number of apparently un­ strom 1974), myctophiforms (Okiyama 1974), ma­ related cottid genera. The genus was considered to rine teleosts in general (Ahlstrom and Moser Constitute a separate family, Hemitripteridae, by 1976), bothids (Futch 1977), scombroids (Okiyama Jordan (1923) and Taranets (1941). and Ueyanagi 1978), serranids (Kendall 1979). In these studies, similarity oflarval form has been in . DISCUSSION remarkable agreement with relationships implied from adult characters. Although larval characters The present state of cottid systematics is con­ have not been used previously as indicators fused and the group and its allies are in need of of relationship (i.e., based on synapomorphies) intensive study (Nelson 1976; Howe and Richard­ among cottids, it seems highly probable that at Son footnote 3). Family limits are not well defined least some of these characters would be as useful (compare, e.g., Jordan 1923; Berg 1940; Taranets in cottids as in other groups offishes. In addition, 1941; Greenwood et al. 1966; Bailey et al. 1970; ifthe cottids were derived from an ancestral stock Nelson 1976). Some genera still need revision and related to the , the most generalized Potential new species remain to be described (e.g., group in the Order (Gill 1889; Nelson 1977; Richardson and Washington 1980; Taranets 1941; Bolin 1947) and if Bolin's (1947) liowe and Richardson footnote 3). Studies of ancestral cottid form is valid and Scorpaenichthys intergeneric relationships have been few (e.g., closely resembles the primitive condition, then Regan 1913; Taranets 1941; Bolin 1947; Watanabe primitive or derived states ofat least some larval 1960) and these had many disagreements (Table characters of cottids can be postulated. Primitive 3). Jordan and Evermann's (1898:1879-1800) com­ states oflarval characters may include four strong Inent of North American Cottidae still has merit, preopercular spines, relatively deep but compact "The family is an extremely varied one which body, compact gut, lack ofgut diverticula, posses­ Cannot be readily thrown into subordinate groups. sion ofa preanal fin fold, rounded snout, relatively ~lrnost every species has an individuality of short pectoral fin. Derived character states may Its own...." include reduced size and/or numbers of preoper­ Because of the confused state of cottid system­ cular spines or modification ofthe basic pattern of atics it seems reasonable to consider whether this four, slender or globose body, trailing gut, pres­ preliminarysummaryof25 generaofcottid larvae ence of gut diverticula, no preanal fin fold, semi­ rnay provide insight into intergeneric relation­ pointed or pointed snout, elongated pectoral fin. ships within the group. Whether these pheneti­ Pigment patterns are more difficult to evaluate as Cally derived larval groupings are indicative of presumably they may possibly reflect responses to relationships among cottid genera depends on habitat or may be more easily modified genetical­ Whether the groups actually possess a set of ly than other morphological characters. This idea shared, derived characters. Determination of de­ has been generally discredited in other groups rived states of larval characters is difficult when where larval characters have been used to imply dealing with such a diverse group as the cottids relationships (e.g., Ahlstrom 1974; Moser and andtheir allies because thelarvaeofmany species Ahlstrom1974; Kendall 1979) as pigmentpatterns are still unknown and complete developmental have substantiated findings based on other char­ Series have not been described for many other acters. Recent experiments on larvae ofthe zebra­ ~pecies. Suchdeterminations arefurther hindered fish, Brachydanio rerio, (Milos and Dingle 1978) 1 the confused state of adult cottid systematics. have demonstrated constancy in numerical regu­ t lthough an in depth analysis of derived charac- lation of melanophores which indicates larval er states is beyond the scope of this study, pigment patterns may not be as plastic as once Consideration of several factors allows discussion thought. Among the cottids, Scorpaenichthys is ~! the ?otential significance of, and possible rela­ heavily pigmented but Enophrys, also considered lonshlpS within, at least some ofthe larval cottid to be a relatively primitive form (Sandercock and groups described here. Larval characters such as Wilimovsky 1968), is not. Heavy pigmentation 117 FISHERY BULLETIN: VOL. 79, NO.1

seems to be related to a neustonic habitat in some based on the distinctiveness of larval characters, (e.g., Scorpaenichthys, Hemilepidotus) but not may warrant consideration at possibly the sub­ others (e.g., Radulinus). Relative constancy of familial level. pigment pattern (such as presence or absence of Group 2 (Paricelinus et al.) shares the derived lateral pigment posterior to the anus) within a slender body form with pointed snout, and also group used in conjunction with other characters, possesses relative constancy ofpigmentation, i.e., however, may provide additional evidence for no lateral pigment. Relationships among at least within-group relationships. some ofthe genera in this grouphave been implied Ifthis line of reasoning and these assumptions previously (Table 3). The distinctiveness oflarval are valid, then certain trends seem apparent form within this group suggests a separate line­ which may be indicative ofrelationships. Group 1 age; this group may warrant possible subfamilial (Artedius et al.) appears to be a natural group status. sharing a number of derived characters not pres­ In Group 3, all but Dasycottus share a highly ent in any other group or genus (Le., multiple modified larval form tending in degrees toward preopercular spines, somewhat trailing gut, un­ globose. The constancy ofthe pigmented pectoral usual gut diverticula, or at least bulging guts). fin is unique among all groups or genera con­ A preanal fin fold is apparently absent and pig­ sidered. With the possible exception ofDasycottus, ment pattern is relatively constant. The grouping the genera appear to bear at least some relation­ agrees with findings of Taranets (1941), in part, ship to each other. and Bolin (1947), who considered the genera to be Group 4 is the most generalized in that a num­ closely related (Table 3). It seems to be a rather ber ofprimitive character states are exhibited and specialized group as Bolin (1947) implied, and, relationships cannot be assessed on given present

TABLE 3.-Intergeneric relationships of cottids as interpreted by A = Regan (1913), B = Taranets (1941), C = Bolin (1947), and D = Watanabe (1960). Included are only those 25 northeast Pacific genera for which larvae are known and discussed in this paper. Parentheses indicate a more distant relationship.

~., ~., ~ ., ~~ ::J ~ ., ~ € '£ .c:.. ~ 0 .c: ~ ~ ~ ~ '" S ~ .!!1'" ~ ., g .~ -., i:!'" s0 8 al- s ~ g. ::J .c: 0 0 ~ g ! .9- 0 0 .c: ~ <: ., .!!1 <: .c: !3 0 ~ .. ~ § ~ <: .. .{;; l'l 0 ~ .g g 8 ~ g. g '" l.Q "" .!!1 ::J .g ~ §' ~ S 8- [ E E ~ -'<'" Cl. ,g> ~ !a ~ .. .c:.. ~ a; ~ ,fi (!l (!l ~ .Q .Q ~ ~ ~ ~ .. .g ~ Genus t3 <3 c "" -.J.. 0 If .r 0: 0: Artedius S/epsias' Chitonotus B.'(C) Ciinocottus B,'C Coitus Dasyeottus Enophrys A Gilbertidie' B Gymnoeanthus' Hemilepidotus (C) (C) Hemitripterus • (A),D JceUnus B,'(C) B,C (C) /ee/us B' B 0 A,B Leptoeottus B C MaJacocottus B,D B Myoxocepha/us B Nautiehthys A,C,D (A),D Ol/gocottus B,'C B,C A A Orthonop/es B,'C B,C B,C Parieel/nus Psychrolutes BB B Radul/nus (C) C (C) C Rhamphocottus 7 AA Seorpaeniehthys· A (C) Trig/ops' A A A 'Considered in family Ble~siidae by Taranets (1941). 'Considered in subfamily Gymnocanthinae by Watanabe (1960). 'Arledius as , useeriops of Taranets (1941). ·Considered in family Hem,tripteridae by Taranets (1941). , Artedius induding Aryias, Allarledius. Astrolytes, 7 Considered in family Rhamphocottidae by Taranets (1941). PeraslTo/rtes of Taranets (1941). 'Considered in order Notothenliformes by Taranets (1941). 'Considered in subfamily Gilbertinae by Watanabe (1960). 'Considered in subfamily Triglopsinae by Watanabe (1960). 118 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE knowledge. Group 5 shares derived character ACKNOWLEDGMENTS states of a very elongate, slender body form and reduced prominence ofpreopercular spines; it also Many who helped make this paper possible were exhibits relative constancy in pigment pattern acknowledged by Richardson and Washington Suggesting relationships between the genera. The (1980). In addition, larvae for illustration were relatively large hatching size (>7 mm NL) oflar­ provided as follows: Dasycottus setiger, J. R. Dunn Vae in Group 6 represents another specialization (Northwest and Alaska Fisheries Center, Nation­ indicative of relationship; such a large size is not al Marine Fisheries Service, NOAA); Psychrolutes known in any ofthe other genera except Hemitrip­ paradoxus, Gilbertidia sigalutes, J. Blackburn terus whose larvae are ca. 12-14 mm NL at (Alaska Department of Fish and Game); ?Mala­ hatching. Group 6 genera share a somewhat cocottus zonurus, P. Wagner and G. Mueller (Uni­ elongate form and constancy ofpigmentation, Le., versity of Alaska); , Myoxo­ lack oflateral pigment, although these characters cephalus polyacanthocephalus, A. Lamb (Pacific alone do not provide strong evidence of relation­ Environment Institute, British Columbia) and ship. That the six ungrouped genera did not share C. Moffett (Bellingham, Wash.). B. Washington a set ofderived characters suggests that they bear illustrated these specimens and provided tech­ no close relationship with one another. nical assistance. N. Y. Khan granted permission In summary, this preliminary examination of to reproduce a figure of Gymnocanthus tricuspis larval characters within 25 genera of cottids has from his dissertation. J. L. Laroche (Oregon State Provided some new insights into cottid systemat­ University) provided information on preopercular ics. Larval' evidence seems to support current spines and pigmentation on Myoxocephalus. Con­ concepts of generic limits in most instances (e.g., versations on cottid systematics with K. Howe Enophrys, Hemitripterus, Hemilepidotus) and (Oregon State University) were particularly in­ has indicated a potentially new northeast Pacific formative and stimulating. K. Howe and B. Wash­ genus represented by "Cottoid Type A." Larval ington read the manuscript and made helpful characters offer support for the distinctiveness of comments. some genera (e.g., Rhamphocottus) and strong relationships among others (e.g., the Artedius LITERATURE CITED group). Some of the larval groupings discussed here tend to support previously implied relation­ AHLSTROM, E. H. ships within the cottids (compare Tables 2 and 3) 1974. The diverse patterns of metamorphosis in gonosto­ but some important differences seem apparent matid fishes - an aid to classification. In J. H. S. Blaxter (editor), The early life history offish, p. 659-674. Springer­ (e.g., the distinctiveness ofGroup 1 demonstrated Verlag, N.Y. herein; theseparation ofArtedius and Icelus, once AHLSTROM, E. H., AND H. G. MOSER. C?nsidered closely related (Jordan 1923); the rela­ 1976. Eggs and larvae offishes and theirrole in systematic tIonship of Paricelinus, generally considered a investigations and in fisheries. Rev. Trav. Inst. Peches Marit.40:379-398. Primitive and rather distinct form (Bolin 1947; BAILEY, R. M., J. E. FITCH, E. S. HERALD, E. A. LACHNER, C. C. Sandercock and Wilimovsky 1968), with other LINDSEY, C. R. ROBINS, AND W B. SCOTT. members ofGroup 2; the apparent relationship of 1970. A list of common and scientific names offishes from [celus to other genera in Group 2 and its question­ the United States and Canada. 3d ed. Am. Fish. Soc. able placement in a separate family (Jordan 1923; Spec. Publ. 6, 149 p. BERG,L.S. Greenwood et al. 1966); the distinctiveness of 1940. A classification offishes, both recent and fossil. Tr. 1?adulinus, previously considered related to Chi­ Zool. Inst. Akad. Nauk SSSR 5(2):87-517. tonotus and Icelinus (Bolin 1947)]. Because of the BERTELSEN, E. Wide diversity of form among cottid larvae, they 1941. The Ceratioid fishes. Ontogeny, , distribu­ tion and biology. Dana-Rep., Carlsberg Found. 39,276 p. offer great potential for helpingto clarify relation­ BIGELOW, H. B., AND W C. SCHROEDER. ships and evolutionary trends within this difficult 1953. Fishes ofthe Gulfof Maine. First Revision. U.S. group of fishes. However, larvae of many species Fish WildL Serv., Fish. Bull. 53, 577 p. remain to be described (rearing may be the best BLACKBURN, J. E. approach), generic limits of larval characters 1973. A survey ofthe abundance, distribution, and factors ~ust affecting distribution of ichthyoplankton in Skagit Bay. be defined, and developmental sequences M.S. Thesis, Univ. Washington, Seattle,136 p. rncluding osteology need to be examined before BOLIN, R. L. that potential can be fully realized. 1941. Embryonic and early larval stages of the cottid 119 FISHERY BULLETIN: VOL. 79, NO.1

fish Orthonopias triads Starks and Mann. Stanford JOHNSON, R. K. Ichthyol. Bull. 2:73-82. 1974. A revision ofthe alepisauroid family Scopelarchidae 1947. The evolution of the marine Cottidae of California (Pisces: Myctophiformes). Fieldiana, Zool. 66, 249 p. with a discussion of the genus as a systematic category. JONES,A. C. Stanford Ichthyo!. Bull. 3:153-168. 1962. The biology of the euryhaline fish Leptocottus BRUUN,A.F. armatus armatus Girard (Cottidae). Univ. Calif. PubI. 1925. On the development and distribution ofthe Norway Zoo!' 67:321-368. bullhead (Cottus lilljeborgi Collett). Cons. Perm. Int. JORDAN, D. S. Explor. Mer, PubI. Circonstance 88, 15 p. 1923. A classification of fishes including families and BUDD, P. L. genera as far as known. Stanford Univ. Publ., Univ. 1940. Development of the eggs and early larvae of six Ser., BioI. Sci. 3:79-243. California fishes. Calif. Div. Fish Game, Fish Bull. 56, JORDAN, D. S., AND B. W. EVERMANN. 50 p. 1898. The fishes of North and Middle America. Bull. COLTON, J. B., JR., AND R. R. MARAK. U.S. Natl. Mus. 47:1241-2183. 1969. Guide for identifying the common planktonic fish KENDALL, A. w., JR. eggs and larvae of continental shelf waters, Cape Sable 1979. Morphological comparisons of North American sea to Block Island. Bur. Commer. Fish., BioI. Lab., Woods bass larvae (Pisces: Serranidae). U.S. Dep. Commer., Hole, Mass., Lab. Ref. 69-9, 43 p. NOAA Tech. Rep. NMFS CIRC-428, 50 p. CUNNINGHAM, J. T. KHAN,N. Y 1891. On some larval stages of fishes. J. Mar. BioI. Assoc. 1972. Comparative morphology and ecology of the pelagic U.K., New Ser., 2:68-74. larvae of nine Cottidae (Pisces) of the northwest Atlantic EHRENBAUM, E. and St. Lawrence drainage. Ph.D. Thesis, Univ. Ottawa, 1904. Eier und Larven von Fischen der deutschen Bucht. 234 p. III. Fische mit festsitzenden Eiern.· Wiss. Meeresunters. KHAN, N. Y., AND D. J. FABER. Komm. wiss. Unters. Dtsch. Meere Kiel BioI. Anst. 1974. A comparison of larvae of the deepwater and Helgo!. 3:127-200. fourhom sculpin, Myoxocephalus quadricornis L. from 1905-9. Eier und Larven von Fischen. Nordisches North America. 1. Morphological development. In J. Plankton, Zool. Teil, 1:1-413. H. S. Blaxter (editor), The early life history of fish, EIGENMANN, C. H. p. 703-712. Springer.Verlag, N.Y. 1892. The fishes of San Diego, California. Proc. U.S. KOEFOED,E. Natl. Mus. 15:123-178. 1907. Poissons. In L. P. R. d'Orleans, Croisiere ocean­ FOLLET'I', W. 1. ographique accompHe a bord de Ill. Belgica dans Ill. Mer 1952. Annotated list of fishes obtained by the California du Gronland 1905, p. 485-499. Charles Bulens, Brussels. Academy of Sciences during six cruises of the U.S.S. KYUSHIN,K. Mulberry conductedbytheUnitedStatesNavy off central 1968. The embryonic development and larval stages California in 1949 and 1950. Proc. Calif. Acad. Sci., of WaBas). BuB. Fac. Fish., Ser. 4, 27:399-432. Hokkaido Univ. 18:277-289. FUIMAN, L. A. 1970. Embryonic development and larvae of Gymno­ 1976. Notes on the early development of the , canthus herzensteini Jordan and Starks. Jpn. J. Ichthyol. Hemitripterusamericanus. Fish. Bull., U.s. 74:467-470. 17:74-79. FUTCH,C.R. LUND, W. A., JR., AND B. C. MARCY, JR. 1977. Larvae of Trichopsetta ventralis (Pisces: Bothiidae), 1975. Early development of the grubby, Myoxocephalus with comments on intergeneric relationships within the aenaeus (MitchiJI). BioI. Bull. (Woods Hole) 149:373-383. Bothidae. Bull. Mar. Sci. 27:740-757. MARLIAVE, J. B. GILL,T. 1975. The behavioral transformation from the planktonic 1889. On the classification of the mail-cheeked fishes. larval stage of some marine fishes reared in the labora­ Proc. U.S. Nat!. Mus. 11:567-592. tory. Ph.D. Thesis, Univ. British Columbia, Vancouver, GoRBUNOVA, N. N. 231 p. 1964. Razmnozhenie i razvitie polucheshuiykh bychkov MCINTOSH, W C., AND A. T. MASTERMAN. (Cottidae, Pisces). (Breeding anddevelopment ofhemilep. 1897. The life-histories of the British marine food-fishes. idotine sculpins (Cottidae, Pisces).) Tr. Inst. Okeanol. C. J. Clay, Lond., 516 p. Akad. Nauk SSSR 73:235·252. [Translated from Russ. MCINTOSH, W C., AND E. E. PRINCE. In T. S. Rass (editor), Fishes of the Pacific and Indian 1890. On the development and life-histories of the tele­ Oceans, biology and distribution, p. 249-266. Isr. Program ostean food- and other fishes. Trans. R. Soc. Edinb. Sci. Transl.,1966, 266 p.; avail. U.S. Dep. Commer., Natl. 35:665-946. Tech. Inf. Serv., Springfield, Va., as 'IT 65-50120.) MILOS, N., AND A. D. DINGLE. GREENWOOD, P. H., D. E. ROSEN, S. H. WEITZMAN, AND 1978. Dynamicsofpigmentpatternformation in thezebra­ G.S.MYERS. fish, Brachydanio reno. II. Liability of lateral line stripe 1966. Phyletic studies of teleostean fishes, with a pro· formation and regulation of pattern defects. J. Exp. visional classification of living forms. Bull. Am. Mus. Zool. 205:217-224. Nat. Rist. 131:341.455. MISITANO, D. A. HAT'I'ORI, S. 1978. Description of laboratory·reared larvae of the buf· 1964. Studies on fish larvae in the Kuroshio and adjacent falo sculpin, Erwphrys bison Girard (Pisces: Cottidae). waters. [In Jpn., Eng!. synop.l Bull. Tokai Reg. Fish. Copeia 1978:635-642. Res. Lab. 40, 158 p. 1980. A record of internal fertilization in the roughback 120 RICHARDSON: CURRENT KNOWLEDGE OF SCULPIN LARVAE

sculpin Chitinotus pugetensis with descriptive notes of RASS, T. S. their early larvae. Copeia 1980:162-165. 1949. The composition of fish fauna of the Barents Sea MORRIS, R. W. and the systematical characters of the fish eggs and 1951. Early development of the cottid fish Clinocottus larvae. [In Russ.] Tr. Vses. Nauchno-Issled. Inst. recalvus (Greeley). Calif. Fish Game 37:281-300. Morsk. Rybn. Khoz. Okeanogr. 17:7-65. MOSER, H. G., AND E. H. AHLSTROM. REGAN, C. T. 1970. Development of lanternfishes (family Myctophidae) 1913. The osteology and classification of the teleostean in the California Current. Part 1. Species with narrow­ fishes of the order Scleroparei. Ann. Mag. Nat. Hist., eyed larvae. Bull. Los Ang. Cty. Mus. Nat. Hist. Sci. ,Ser. 8, 11:169-184. 7,145 p. RICHARDSON, S. L., AND B. B. WASHINGTON. 1972. Development of the lanternfish, Scopelopsis multi­ 1980. Guide to identification of some sculpin (Cottidae) punctatus Brauer 1906, with a discussion of its phylo­ larvae from marine and brackish waters off Oregon and genetic position in the family Myctophidae and its role adjacent areas of the northeast Pacific. U.S. Dep. in a proposed mechanism for the evolution of photophore Commer., NOAA Tech. Rep. NMFS CIRC-430, 56 p. patterns in lanternfishes. Fish. Bull., U.S. 70:541-564. RUSSELL, F. S. 1974. Role of larval stages in systematic investigations of 1976. The eggs and planktonic stages of British marine marine teleosts: The Myctophidae, a case study. Fish. fishes. Acad. Press, Lond., 524 p. Bull., U.S. 72:391-413. NELSON, J. S. SANDERCOCK, F. K., AND N. J. WILIMOVSKY. 1968. Revision of the cottid genus Enophrys. Copeia 1976. Fishes of the world. Wiley, N.Y., 416 p. 1968:832-853. 1977. Fishes ofthe southern hemisphere genus Neophryn­ ichthys (Scorpaenifonnes: Cottoidei), with descriptions of STEIN, R. two new species from New Zealand and Macquarie Island. 1972. Identification of some larval Pacific cottids. M.S. J. R. Soc. N.z. 7:485-511. Thesis, Humboldt State Coli., Arcata, Calif., 41 p. O'CONNELL, C. P. 1973. Description of laboratory-reared larvae of Oligocot­ 1953. The life history of the Scorpaenichthys tus maculosus Girard (Pisces:Cottidae). Copeia 1973: marmoratus (Ayres). Calif. Dep. Fish Game, Fish Bull. 373-377. 93,76 p. TARA NETS, A. Y. OKIYAMA,M. 1941. K klassifikatsii iproiskhozhdenifbychkovsemeystva 1974. The larval taxonomy ofthe primitive myctophifonn Cottidae. (On the classification and origin of the family fishes. In J. H. S. Blaxter (editor), The early life history Cottidae.) Izvestiya Akad. Nauk SSSR. Otd. Bio!. 1941 offish, p. 609-621. Springer-Verlag, N.Y. (3):427-447. [Translated from Russ., Inst. Fish., Univ. OKIYAMA, M., AND H. SANDO. B.C., Mus. Contrib. 5, 28 p.) 1976. Early life history of the sea raven, Hemitripterus WARFEL, H. E., AND D. MERRIMAN. villosus, (Hemitripterinae, Cottidae) in the Japan Sea. 1944. The spawning habits, eggs, and larvae of the sea [In Jpn., Eng!. abstr.] Bull. Jpn. Sea Reg. Fish. Res. raven, Hemitripterus americanus, in southern New Lab. 27:1-10. England. Copeia 1944:197-205. O!{JYAMA, M., AND S. UEYANAGI. 1978. Interrelationships of scomhroid fishes: an aspect WATANABE, M. from larval morphology. Bull. Far Seas Fish. Res. Lab. 1960. Fauna Japonica Cottidae (Pisces). Tokyo News (Shimizu) 16:103-113. Service, 218 p. PEDEN,A. E. WHITE, W. A. 1978. A systematic revision of the hemilepidotine fishes 1977. Taxonomic composition, abundance, distribution (Cottidae). Syesis 11:11-49. and seasonality of fish eggs and larvae in Newport Bay, PERLMUTI'ER, A. California. M.S. Thesis, California State Univ., fuller­ 1939. Section 1. An ecological survey of young fish and ton, 107 p. eggs identified from tow-net collections. In A biological ZVJAGINA, O. A. survey of the salt waters of Long Island, 1938, Part II, 1963. Materialy po razmnozheniyu i razvituju ryb morya p. 11-71. N.Y. Conserv. Dep., Supp!. 28th Annu. Rep., Laptevykh 2. Ledovitomorskaya rogatka, i 3. Aziatskaya 1938, Salt-water Surv. 15. Koryushka. (Materials on the reproduction and develop­ QUAST,J. C. ' ment of fish of the Laptev Sea 2. Arctic sculpin and 1965. Osteological characteristics and affinities of the 3. Asiatic smelt.) [In Russ., Eng!. summ.] Tr. Inst. hexagrammid fishes, with a synopsis. Proc. Calif. Acad. Okeanol. 62:3-12. [Eng!. trans!. in Lib. Nat. Mus. Can., Sci., Ser. 4, 31:563-600. Ottawa.]

121