Description of early life history stages of the northern sculpin (Icelinus borealis Gilbert) (Teleostei: Cottidae).Abstract--Larvae of the genus Icelinus are collected more frequently than any other sculpin sculpin, common name for a member of the large family Cottidae, bizarre fishes with large, spiny or armored heads and short, tapering bodies, found in both marine and freshwater habitats. The family includes the muddlers and some species called bullheads. larvae Larvae, in Roman religion Larvae: see lemures. in ichthyoplankton surveys in the Gulf of Alaska Noun 1. Gulf of Alaska - a gulf of the Pacific Ocean between the Alaska Peninsula and the Alexander Archipelago Pacific, Pacific Ocean - the largest ocean in the world and Bering Sea Bering Sea, c.878,000 sq mi (2,274,020 sq km), northward extension of the Pacific Ocean between Siberia and Alaska. It is screened from the Pacific proper by the Aleutian Islands. The Bering Strait connects it with the Arctic Ocean. , and larvae of the northern sculpin (Icelinus borealis) are commonly found in the ichthyofauna ich·thy·o·fau·na n. The fish of a particular region. in both regions. Northern sculpin are geographically isolated north of the Aleutian Islands Aleutian Islands (əl  `shən), chain of rugged, volcanic islands curving c.1,200 mi (1,900 km) west from the tip of the Alaska Peninsula and approaching Russia's Komandorski Islands. , Alaska, which allows for a definitive description of
its early life history development in the Bering Sea. A combination of
morphological characters, pigmentation pigmentation, name for the coloring matter found in certain plant and animal cells and for the color produced thereby. Pigmentation occurs in nearly all living organisms. , preopercular spine pattern,
meristic counts, and squamation in later developmental stages is
essential to identify Icelinus to the species level. Larvae of northern
sculpin have 35-36 myomeres, pelvic fins with one spine and two rays, a
bony preopercular shelf, four preopercular spines, 3-14 irregular
postanal ventral ventral /ven·tral/ (ven´tral)1. pertaining to the abdomen or to any venter. 2. directed toward or situated on the belly surface; opposite of dorsal. ven·tral adj. melanophores, few, if any, melanophores ventrally on the gut, and in larger specimens, two rows of ctenoid scales directly beneath the dorsal fins extending onto the caudal caudal /cau·dal/ (kaw´d'l) 1. pertaining to a cauda. 2. situated more toward the cauda, or tail, than some specified reference point; toward the inferior (in humans) or posterior (in animals) end of the body. peduncle peduncle /pe·dun·cle/ (pe-dung´k'l) a stemlike connecting part, especially (a) a collection of nerve fibers coursing between different areas in the central nervous system, or (b) . The taxonomic characters of the larvae of northern sculpin in this study may help differentiate northern sculpin larvae from its congeners, and other sympatric sym·pat·ric adj. Ecology Occupying the same or overlapping geographic areas without interbreeding. Used of populations of closely related species. sculpin larvae, and further aid in solving complex systematic relationships within the family Cottidae. ********** The sculpin family, Cottidae, is a speciose, morphologically diverse group of fishes with a worldwide distribution comprising as many as 275 species in about 70 genera (Nelson, 2006). Greatest diversity occurs in the Northeast Pacific Ocean and Bering Sea with 96 species in 34 genera where they are found in almost every benthic ben·thos n. 1. The collection of organisms living on or in sea or lake bottoms. 2. The bottom of a sea or lake. [Greek. habitat from the intertidal in·ter·tid·al adj. Of or being the region between the high tide mark and the low tide mark. in to the upper continental slope (Mecklenburg et al., 2002; Nelson, 2006; Pietsch and Orr, 2006). Cottids are primarily predators of smaller fish and crustaceans, and many species are preyed upon by larger fishes and marine mammals, particularly pinnipeds (Browne et al., 2002; Pietsch and Orr, 2006). Cottids are one of several prey species exploited by the harbor seal harbor seal, most commonly seen seal of the Northern Hemisphere, Phoca vitulina. Harbor seals are found along coasts and in sheltered bays and harbors of North America, Europe, and NE Asia. (Phoca vitulina Phoca vitulina see harbor seal. ). Cottid species, including the northern sculpin (Icelinus borealis), are abundant in waters surrounding rookeries of Steller sea lions (Eumetopias jubatus) where they contribute to the diversity of available prey species (Mueter and Norcross, 2000; Browne et al., 2002; Fritz and Hinckley, 2005). New cottid species continue to be described; however, the systematics systematics: see classification. and life histories of most species are poorly known. A more complete understanding of the diversity of the family is necessary to fully understand their role in the dynamics of North Pacific ecosystems (Hoff, 2006; Pietsch and Orr, 2006). Icelinus borealis is the most common species of Icelinus in the Gulf of Alaska and the only species of Icelinus known from the Bering Sea. It is reported to be an important component of the ichthyofauna in both regions (Mueter and Norcross, 2000; Mecklenburg et al., 2002). Adults are distributed from Attu Island in the Aleutian Islands and Bristol Bay in the eastern Bering Sea to southern Puget Sound, Washington, at depths of 4-247 m, on nearly all types of substrate (Mecklenburg et al., 2002). Larvae of Icelinus are the most frequently collected larval larval 1. pertaining to larvae. 2. larvate. larval migrans see cutaneous and visceral larva migrans. cottids in the Northeast Pacific Ocean and Bering Sea, occurring in 9.3% (ranked 12th of all taxa collected) of ichthyoplankton samples collected by the Alaska Fisheries Science Center (AFSC AFSC American Friends Service Committee AFSC Alaska Fisheries Science Center AFSC Air Force Systems Command AFSC Air Force Specialty Code AFSC Air Force Space Command AFSC Armed Forces Services Corporation AFSC Army Field Support Command ). Larvae of Icelinus have primarily been collected in continental shelf and slope waters of the Bering Sea, through Unimak Pass to the Gulf of Alaska and Shelikof Sea Valley, around Kodiak Island, and southward to the west coast of the United States The "West Coast", "Western Seaboard", or "Pacific Seaboard" are terms for the westernmost coastal states of the Western United States, comprising most often California, Oregon and Washington. . In the Shelikof Sea Valley, they are most often collected along the northern side, closest to the Alaska Peninsula (Matarese et al., 2003). Icelinus comprises 11 species that are diagnosed by pelvic fins having one spine and two rays, four preopercular spines (the dorsalmost is longest and bifid bifid /bi·fid/ (bi´fid) cleft into two parts or branches. bi·fid adj. Forked or split into two parts. or trifid trifid /tri·fid/ (tri´fid) split into three parts. tri·fid adj. Divided into three narrow parts or lobes. trifid split into three parts. ), two rows of ctenoid scales directly beneath the dorsal fins, and gill membranes that are united and free from the isthmus isthmus (ĭs`məs), narrow neck of land connecting two larger land areas. Since it commands the only land route between two large areas and is on two seas, an isthmus has great strategical and commercial importance and is a favorable situation (Bolin, 1936; Yabe et al., 1983; Yabe et al., 2001; Nelson et al., 2004; Rosenblatt and Smith, 2004). Adult I. borealis reach 10 cm standard length and lack distinct postocular spines, possess a long cirrus at the base of the nasal spine, the first or second dorsal-fin spines are not longer than the third or fourth, and the two rows of ctenoid scales below the dorsal fins extend onto the caudal peduncle (Bolin, 1936; Mecklenburg et al., 2002). This study is the first to identify and describe the larval and juvenile stages of I. borealis. Previous descriptions were based on misidentified specimens or were made at a more conservative generic level because of difficulty distinguishing among species of Icelinus and between Icelinus and other sympatric cottid larvae. Larvae of Icelinus quadriseriatus from the coast of California are currently the only Icelinus larvae described (Feeney, 1987). Larvae tentatively identified as I. borealis in early literature were misidentified as Ruscarius meanyi based on a pelvic-fin ray count of 1, 2--a count diagnostic of Icelinus but also occurring rarely in R. meanyi (Blackburn, 1973; Richardson, 1977; Richardson and Pearcy, 1977; Richardson and Washington, 1980; Washington, 1981; Begle, 1989). Current literature has continued to identify larvae of Icelinus at the generic level; however, Matarese et al. (1989, 2003) have cautiously identified illustrations as I. borealis. Icelinus borealis has cautiously been identified at the species level because three other species of Icelinus with unidentified larvae (I. burchami, L filamentosus, and I. tenuis ten·u·is n. pl. ten·u·es Linguistics 1. A voiceless stop. 2. A voiceless unaspirated stop in ancient Greek. ), and other unidentified cottid larvae (e.g., Icelus) are also found in the Gulf of Alaska. Uniformity between larval Icelinus and other cottid larvae is noted in the assignment of phenetic phe·net·ic adj. Of, relating to, or designating a system of classification of organisms based on overall or observable similarities rather than on phylogenetic or evolutionary relationships. groups based on shared larval characters (e.g., preopercular spine pattern, body shape, and pigmentation) (Richardson, 1981). Icelinus is included in phenetic group 2, which includes Paricelinus, Triglops, Icelus (tentatively), and Chitonotus, and is characterized by a slender body shape, pointed snout snout the upper lip and the apex of the nose, especially of the pig. Called also rostrum. Has a specialized skin to survive the rigors of rooting, is supported by a separate bone (the os rostri), and also has a few sensory hairs. , and four prominent preopercular spines (Richardson, 1981). Further study of phenetic groups has increased the size of group 2, the "Myoxocephalus group," to include a total of 13 genera (Matarese et al., 1989; Moser et al., 1996). The Myoxocephalus group includes the genera previously included in group 2 as well as the genera Myoxocephalus, Ruscarius, Ascelichthys, Orthonopias, Enophrys, Radulinus, Gymnocanthus, and Synchirus (Matarese et al., 1989; Moser et al., 1996). Members of the Myoxocephalus group have four preopercular spines and are defined by a unique larval character, namely a bony preopercular shelf (Moser et al., 1996). Larvae of Icelinus are reported to be the most frequently collected larval cottids in the Northeast Pacific Ocean and Bering Sea. Although collected in large numbers, the size range of specimens is limited, which has hindered compiling the developmental series necessary for description. Increased ichthyoplankton sampling conducted in the Bering Sea in the 1990s has provided the specimens necessary to describe larvae of I. borealis using meristic counts and morphological characters, including pigmentation and preopercular spination. This study presents an illustrated developmental series and general aspects of osteological development for L borealis. Methods A total of 53 specimens (7.4-51.7 mm standard length [SL]) collected during AFSC research cruises in the Bering Sea and Gulf of Alaska between 1979 and 2002 were examined (Fig. 1). Specimens were collected at depths to 400 m, primarily using 60-cm bongo bongo (bŏng`gō), spiral-horned antelope, Boocercus eurycerus, found in jungles and thick bamboo forests of equatorial Africa. Shy, elusive animals, bongos never emerge into the open and are seldom seen; they browse singly or in small nets and Methot trawls. Specimens were initially preserved in 5% formalin formalin /for·ma·lin/ (for´mah-lin) formaldehyde solution. for·ma·lin n. An aqueous solution of formaldehyde that is 37 percent by weight. buffered with sodium borate sodium borate n. A crystalline compound that is the sodium salt of boric acid and is used as an alkalizing agent and as a mild astringent in lotions, gargles, and mouthwashes. , then later transferred to 70% ethanol. Nineteen specimens were cleared and stained using the method of Potthoff (1984). Twenty-two adult Icelinus borealis specimens were radiographed to verify the vertebral ver·te·bral adj. 1. Of, relating to, or of the nature of a vertebra. 2. Having or consisting of vertebrae. 3. Having a spinal column. count of 35-36 recorded in literature. Specimens were grouped using the series method, by positively identifying juveniles using known adult characteristics then linking those specimens to progressively smaller specimens using shared characteristics (Neira et al., 1998). Larvae were identified using reported generic characters for Icelinus including 35-36 vertebrae Vertebrae Bones in the cervical, thoracic, and lumbar regions of the body that make up the vertebral column. Vertebrae have a central foramen (hole), and their superposition makes up the vertebral canal that encloses the spinal cord. (myomeres) and four distinct preopercular spines, if developed. Illustrated Icelinus (tentatively identified as I. borealis) from Matarese et al. (1989) were also used to compare general morphological and pigment characters. Meristic counts are reported for ossified os·si·fy v. os·si·fied, os·si·fy·ing, os·si·fies v.intr. 1. To change into bone; become bony. 2. elements using cleared and stained or radiographed material. Morphometric measurements were taken following Richardson and Washington (1980) using a digital image analysis system with Image Pro Plus, vers. 4.5 software (Media Cybernetics, Inc., Silver Spring, MD). Both body length and proportional measurements are in SL unless otherwise noted. Developmental terminology follows Kendall et al. (1984). Nomenclature describing caudal-fin development follows Matarese and Marliave (1982). Only melanistic mel·a·nism n. 1. See melanosis. 2. Dark coloration of the skin, hair, fur, or feathers because of a high concentration of melanin. mel pigmentation is described. Nomenclature describing pigment pattern follows Busby and Ambrose (1993). The term "band" refers to an aggregation of melanophores oriented vertically; "bar" refers to an aggregation that is oriented horizontally. Illustrations were rendered using a camera lucida attached to a dissecting dis·sect tr.v. dis·sect·ed, dis·sect·ing, dis·sects 1. To cut apart or separate (tissue), especially for anatomical study. 2. stereomicroscope ster·e·o·mi·cro·scope n. A microscope equipped for stereoscopic viewing. ster  e·o·mi .
Material examined Larvae: 53 specimens examined, 7.4-51.7 mm. UW 105110, 1 (16.7 mm), Bering Sea, 52[degrees]35.9'N, 173[degrees]25.6'W, 137 m depth, 2 August 1997, FV Vesteraalen; UW 105111, 1 (13.4 mm), Bering Sea, 56[degrees]31.9'N, 166[degrees]25.4'W, 88 m depth, 16 July 1994, RV Miller Freeman; UW 105113, 2 (14.4-15.1 mm), Bering Sea, 56[degrees]30.6'N, 168[degrees]60.0'W, 95 m depth, 23 July 2001, TS Oshoro maru; UW 105114, 1 (12.1 mm), Bering Sea, 54[degrees]59.7'N, 166[degrees]58.9'W, 100 m depth, 19 July 1995, TS Oshoro maru; UW 105116, 1 (14.5 mm), Bering Sea, 56[degrees]59.6'N, 170[degrees]00.4'W, 62 m depth, 25 July 1996, TS Oshoro maru; UW 105117, 2 (13.4-14.3 mm), Bering Sea, 57[degrees]01.1'N, 171[degrees]00.2'W, 94 m depth, 25 July 1996, TS Oshoro maru; UW 105119, 2 (14.3-16.3 mm), Bering Sea, 55[degrees]00.9'N, 166[degrees]01.4'W, 100 m depth, 21 July 1997, TS Oshoro maru; UW 105121, 1 (32.1 mm), Gulf of Alaska, 58[degrees]12.1'N, 150[degrees]27.0'W, 115 m depth, 16 May 1985, RV Poseidon; UW 105122, 1 (7.9 mm), Bering Sea, 54[degrees]01.3'N, 166[degrees]33.9'W, 100 m depth, 25 April 1993, RV Miller Freeman; UW 105124, 1 (51.7 mm), Gulf of Alaska, 55[degrees]55.5'N, 157[degrees]56.0'W, 94 m depth, 23 June 1998, RV Wecoma; UW 105125, 2 (10.7-15.2 mm), Gulf of Alaska, 58[degrees]22.1'N, 151[degrees]22.2'W, 100 m depth, 1 June 2002, RV Miller Freeman; UW 105127, 1 (8.3 mm), Bering Sea, 54[degrees]55.5'N, 165[degrees]29.1'W, 119 m depth, 23 May 2003, RV Miller Freeman; UW 105129, 2 (9.2-9.3 mm), Bering Sea, 56[degrees]27.3'N, 169[degrees]28.3'W, 94 m depth, 12 July 1997, RV Wecoma; UW 105131, 1 (8.8 mm), Bering Sea, 56[degrees]27.3'N, 169[degrees]28.3'W, 30 m depth, 12 July 1997, RV Wecoma; UW 105133, 2 (8.5-12.5 mm), Bering Sea, 56[degrees]30.2'N, 169[degrees]28.5'W, 78 m depth, 10 July 1997, RV Wecoma; UW 105134, 1 (14.9 mm), Bering Sea, 56[degrees]41.4'N, 169[degrees]48.5'W, 74 m depth, 8 July 1997, RV Wecoma; UW 105136, 2 (8.0-9.2 mm), Bering Sea, 56[degrees]42.6'N, 169[degrees]35.9'W, 64 m depth, 10 July 1997, RV Wecoma; UW 105138, 1 (13.2 mm), Bering Sea, 56[degrees]42.6'N, 169[degrees]35.9'W, 25 m depth, 10 July 1997, RV Wecoma; UW 105140, 2 (10.0-14.1 mm), Bering Sea, 56[degrees]42.6'N, 169[degrees]36.1'W, 70 m depth, 9 July 1997, RV Wecoma; UW 105142, 1 (14.9 mm), Bering Sea, 56[degrees]42.7'N, 169[degrees]36.5'W, 72 m depth, 9 July 1997, RV Wecoma; UW 105144, 1 (10.2 mm), Bering Sea, 56[degrees]53.2'N, 170[degrees]26.7'W, 87 m depth, 6 July 1997, RV Wecoma; UW 105146, 2 (15.4-16.5 mm), Bering Sea, 57[degrees]17.3'N, 170[degrees]10.1'W, 30 m depth, 6 July 1997, RV Wecoma; UW 105148, 1 (14.5 mm), Bering Sea, 57[degrees]21.2'N, 170[degrees]08.3'W, 50 m depth, 13 July 1997, RV Wecoma; UW 105149, 1 (7.4 mm), Bering Sea, 54[degrees]24.9'N, 165[degrees]09.0'W, 140 m depth, 25 April 1997, RV Miller Freeman; UW 105151, 3 (43.2-45.9 mm), Gulf of Alaska, 57[degrees]18.5'N, 152[degrees]02.8'W, 74 m depth, 13 September 1993, RV Miller Freeman; UW 105152, 1 (41.7 mm), Gulf of Alaska, 57[degrees]15.7'N, 152[degrees]53.7'W, 87 m depth, 16 September 1993, RV Miller Freeman; UW 105154, 1 (14.1 mm), Bering Sea, 56[degrees]32.0'N, 166[degrees]25.4'W, 88 m depth, 16 July 1994, RV Miller Freeman; UW 105156, 1 (19.6 mm), Bering Sea, 57[degrees]24.9'N, 170[degrees]05.6'W, 52 m depth, 13 September 1999, RV Miller Freeman; UW 105157, 1 (11.6 mm), Gulf of Alaska, 56[degrees]46.2'N, 156[degrees]46.7'W, 101 m depth, 27 May 1995, RV Miller Freeman; UW 105159, 1 (9.4 mm), Gulf of Alaska, 57[degrees]24.5'N, 155[degrees]48.6'W, 100 m depth, 28 May 1995, RV Miller Freeman; UW 105160, 1 (17.9 mm), Bering Sea, 55[degrees]04.4'N, 165[degrees]08.0'W, 108 m depth, 26 July 1996, RV Miller Freeman; UW 105162, 2 (13.4-15.8 mm), Bering Sea, 56[degrees]28.3'N, 169[degrees]26.9'W, 87 m depth, 1 August 1996, RV Miller Freeman; UW 105164, 2 (11.1-12.9 mm), Bering Sea, 56[degrees]30.3'N, 171[degrees]02.5'W, 119 m depth, 4 August 1996, RV Miller Freeman; UW 105165, 3 (14.8-16.0 mm), Bering Sea, 56[degrees]32.7'N, 169[degrees]27.4'W, 63 m depth, 2 August 1996, RV Miller Freeman; UW 105167, 1 (13.8 mm), Bering Sea, 56[degrees]34.6'N, 169[degrees]24.3'W, 44 m depth, 2 August 1996, RV Miller Freeman; UW 105169, 1 (24.9 mm), Bering Sea, 56[degrees]31.2'N, 169[degrees]28.8'W, 68 m depth, 11 September 1997, RV Miller Freeman; UW 105172, 1 (24.1 mm), Bering Sea, 57[degrees]17.3'N, 170[degrees]09.3'W, 39 m depth, 16 September 1997, RV Miller Freeman; UW 105174, 1 (22.7 mm), Bering Sea, 57[degrees]16.3'N, 170[degrees]11.0'W, 16 September 1997, RV Miller Freeman. [FIGURE 1 OMITTED] Adults: 22 specimens examined, 32.0-77.0 mm. UW 027383, 4 (41.0-50.0 mm), eastern North Pacific, 60[degrees]12.0'N, 147[degrees]45.0'W, 30 m depth, 1 August 1989, RV Discovery, J. W. Orr; UW 029499, 5 (32.0-55.0 mm), eastern North Pacific, 60[degrees]21.0'N, 147[degrees]49.0'W, 40 m depth, 6 August 1989, RV Discovery, J. W. Orr; UW 040432, 3 (45.0-64.0 mm), eastern North Pacific, 60[degrees]18.0'N, 147[degrees]50.0'W, 142 m depth, 31 July 1989, RV Discovery, C. Eaton; UW 111416, 2 (55.0-62.0 mm), eastern North Pacific, 52[degrees]39.8'N, 169[degrees]21.6'W, 114 m depth, 24 May 2003, FV Northwest Explorer, J. W. Orr; UW 040955, 4 (44.0-45.0 mm), eastern North Pacific, 60[degrees]33.2'N, 147[degrees]35.0'W, 40 m depth, 1 October 1989, A. M. Shedlock; UW 027174, 4 (60.0-77.0 mm), eastern North Pacific, Gulf of Alaska, Yakutat Bay, FV Resolution. Results Morphology The smallest larva larva, in zoology larva, independent, immature animal that undergoes a profound change, or metamorphosis, to assume the typical adult form. Larvae occur in almost all of the animal phyla; because most are tiny or microscopic, they are rarely seen. examined in this study was 7.4 mm notochord notochord (nō`təkôrd'), in biology, supporting rod running most of the length of animals of the phylum Chordata and present at varying times in the life cycle. length (NL) and in preflexion (Fig. 2A). Notochord flexion flexion /flex·ion/ (flek´shun) the act of bending or the condition of being bent. flex·ion n. 1. The act of bending a joint or limb in the body by the action of flexors. 2. began at approximately 8.0 mm and was complete around 11.0 mm (Fig. 2B). Postflexion larvae were 11.0-16.0 mm (Fig. 2C). Transformation to the juvenile stage occurred between 16.0 mm and 24.0 mm (Fig. 2D). Specimens larger than 24.0 mm were considered juveniles and identified using adult characters (Fig. 2E). During preflexion, the head was small and round, measuring 18% SL, increasing to 38% SL by the juvenile stage (Table 1). The snout was initially rounded, but became notably pointed by flexion; snout length was 24% head length (HL) during preflexion, increasing to approximately 28% HL during flexion through the juvenile stage. Snout-to-anus length steadily increased from 39% SL during preflexion to 51% SL in juveniles. Body depth was initially 17% SL during preflexion, but increased to approximately 20% SL in later stages. Pigmentation Two preflexion specimens were available for study: one 7.4 mm NL and one 7.9 mm NL. Both specimens were lightly pigmented (Fig. 2A). A single melanophore melanophore /mel·a·no·phore/ (-for?) a pigment cell containing melanin, especially such a cell in fishes, amphibians, and reptiles. mel·a·no·phore n. was present on the lower jaw angle. Pigment on the gut consisted of one to three individual melanophores anteriorly, and moderate pigmentation on the anus. A single row of nine postanal ventral melanophores (PVMs) was present on both specimens. Pigmentation on the caudal finfold was present on the 7.4 mm NL specimen. Pigment on the head, gut, and anus steadily increased during flexion (Fig. 2B). Twenty-six postflexion and transforming specimens were examined. Melanophores were present dorsally over the mid- and hind-brain (Fig. 2C). Minute melanophores were present on the orbital rim. Loosely grouped melanophores were present in postorbital and suborbital suborbital /sub·or·bi·tal/ (sub-or´bi-t'l) infraorbital. sub·or·bit·al adj. Situated on or below the floor of the orbit of the eye. n. regions, upper and lower jaws, on the cheek, operculum operculum /oper·cu·lum/ (o-per´ku-lum) pl. oper´cula [L.] 1. a lid or covering. 2. the folds of pallium from the frontal, parietal, and temporal lobes of the cerebrum overlying the insula. , chin, and isthmus. Pigment was present on the nape. The gut was pigmented along the anterodorsal surface and extended dorsolaterally toward the anus. Three to 14 PVMs were present on specimens between preflexion and postflexion stages; nine was the modal value (Table 2). The size, shape, and location of PVMs were variable among specimens. Lateral body pigment developed as vertical (dorsal to ventral) bands that were composed of densely aggregated small melanophores. The anterior (first) lateral band was located directly under the first dorsal fin and extended ventrally to the gut. The second band developed as a small aggregation of melanophores located mediolaterally on the body. When fully developed, the second band extended from the anterior portion of the second dorsal fin to the mediolateral part of the body. Pigment developed on the first dorsal fin, particularly on the membrane between the first two or three spines. Rays of the second dorsal fin were also pigmented in some specimens. Large, dark melanophores were present on the pectoral-fin base, and some pigmentation developed on the rays near the base. One or two melanophores were present on or near the pelvic-fin base. [FIGURE 2 OMITTED] Throughout transformation of larvae of L borealis into the juvenile stage, pigmentation continued to increase on the head until the entire area was nearly covered with small melanophores (Fig. 2D). Gut pigment was less visible. The first and second lateral pigment bands were fully developed. The third lateral pigment band developed directly beneath the posterior portion of the second dorsal fin approximately between fin rays 11 and 13. Pigment in the fourth band was located on the caudal peduncle and extended posteriorly onto the caudal fin. Pigment was also scattered mediolaterally, giving the appearance of a horizontal bar posterior to the second band. Pigment developed on the caudal-fin rays. At the beginning of the juvenile stage, lateral bands were well defined by dark pigment (Fig. 2E). Larval pigmentation (e.g., PVMs) was still present until at least 24.9 mm, but by 33.0 mm no residual larval pigment remained. Scattered melanophores on the mediolateral part of the body between the second and fourth bands were retained and looked like small pigment blotches. Cirri cir·ri n. Plural of cirrus. Cirri developed during the postflexion stage. Supraocular cirri were first to develop. Supraocular cirri are typically bifid or trifid, but occasionally have more than three terminal filaments. The development of nasal and postorbital cirri followed supraocular cirri. During transformation into the juvenile stage, cirri developed posterodorsally on the maxilla maxilla /max·il·la/ (mak-sil´ah) pl. maxil´las, maxil´lae [L.] the irregularly shaped bone that with its fellow forms the upper jaw. max´illary max·il·la n. pl. . By 25.0 mm, one small cirrus was present both anteriorly and posteriorly of the parietal parietal /pa·ri·e·tal/ (pah-ri´e-t'l) 1. of or pertaining to the walls of a cavity. 2. pertaining to or located near the parietal bone. pa·ri·e·tal adj. 1. and nuchal spines, and more than one opercular o·per·cu·lum n. pl. o·per·cu·la or o·per·cu·lums A lid or flap covering an aperture, such as the gill cover in some fishes or the horny shell cover in snails or other mollusks. cirrus may develop per side (two cirri were present dorsally on each opercle of a 46.0-mm specimen). A full complement of supraocular, nasal, postorbital, maxillary max·il·lar·y adj. Of or relating to a jaw or jawbone, especially the upper one. n. A maxillar; a jawbone. maxillary (mak´siler´ē), adj , occipital, and opercular cirri was present in juveniles. Meristic features Except for the dorsal-fin spines and rays and the superior procurrent caudal-fin rays, fins ossified by 14.3 mm (Table 3). Dorsal-fin spines and rays were completely ossified at 15.8 mm, as were the superior procurrent caudal-fin rays. Vertebral centra (9-11 abdominal + 24-27 caudal) ossified at 14.3-15.8 mm. By 15.0 mm, lateral line scales began to develop; by 15.8 mm, two dorsal scale rows began to develop immediately beneath the dorsal fins. Lateral line scales and the two dorsal scale rows were ossified by 24.0 mm. Pterygiophores of the dorsal and anal fins ossified by 24.1 mm. Adult radiographs resulted in vertebral counts of 35-36. Spination Head spines developed during flexion. At 8.8 mm, parietal spines were minute but ossified. Four preopercular spines were present; the dorsalmost spine was most pronounced. At 11.6 mm, small nuchal spines, approximately half the size of the parietals, were present. By 14.3 mm, nasal spines were ossified. Parietal and nuchal spines fused together at their tips to form parietal sensory canals. By 16.0 mm, the dorsalmost preopercular spine was bent upward and the ventralmost spine downward and forward. The fused parietal and nuchal spines were less prominent. Nasal spines were well developed and slightly curved posteriorly by 22.7 mm. At approximately 24.0 mm, the dorsalmost preopercular spine was very large and bifurcate To divide into two. ; the dorsalmost spine may become trifurcate by the juvenile stage. Caudal skeleton The caudal skeleton consisted of one ural centrum centrum /cen·trum/ (sen´trum) pl. cen´tra [L.] 1. a center. 2. the body of a vertebra. cen·trum n. pl. cen·trums or cen·tra 1. , preural centra, neural and haemal spines, three epurals, two uroneurals, one superior hypural ([HY.sub.4.5]), one inferior hypural ([HY.sub.1-3]), and 25-31 caudal-fin rays (7-11, 6 + 6, 4-8) (Fig. 3). At 8.8 mm, [HY.sub.1-3] and [HY.sub.4-5] were fused and all 12 principal caudal-fin rays (6 + 6) were present (Fig. 3A). Three epurals formed by 12.0 mm. Each preural centrum had one neural and one haemal spine; however, in some specimens the first preural centrum had two neural spines (Fig. 3B). All five hypurals ([HY.sub.1-5]) fused by 13.4 mm. By 15.8 mm, the ural centrum, preural centra, and principal caudal-fin rays ossified (Fig. 3C). Hypurals ossified by 16.0 mm. Two uroneurals were present and ossified by 20.0 mm; neural and haemal spines on the first preural centrum and procurrent caudal-fin rays were ossified. Epurals ossified by 22.7 mm (Fig. 3D). By the juvenile stage at approximately 24.0 mm, development of the caudal skeleton was complete. [FIGURE 3 OMITTED] Discussion Information about the early life history of Icelinus is conspicuously sparse in literature. This study presents the first description of larval and juvenile Icelinus borealis. Icelinus borealis larvae exhibit a unique geographic distribution in the Bering Sea and are geographically isolated north of the Aleutian Islands--which provides for a definitive description of its development. A combination of morphological characters, pigmentation, preopercular spine pattern, meristic counts, and squamation in later developmental stages is essential to identify Icelinus at the species level. Larvae of I. borealis have 35-36 myomeres. The body is lightly pigmented, and the most useful character is the presence of 3-14 (mode = 9) irregular PVMs that persist through transformation into the juvenile stage. Four prominent preopercular spines and three rows of spiny spiny sharp spines protrude. spiny amaranth amaranthusspinosum. spiny anteater see echidna. spiny clotburr xanthiumspinosum. spiny emex see emex australis. ctenoid scales develop during transformation into the juvenile stage; one row is along the lateral line and two are directly beneath the dorsal fins. Identification of I. borealis larvae in other geographic areas, such as the Gulf of Alaska, is complicated by the co-occurrence of other species of Icelinus. Icelinus filamentosus is found with I. borealis throughout the Gulf of Alaska but, if collected, has not been identified in ichthyoplankton samples (Matarese et al., 1989; Mecklenburg et al., 2002). Larvae of I. borealis differ from I. filamentosus primarily by having an anal-fin ray count of 11-14 (vs. 13-16) and a vertebral count of 35-36 (vs. 34-37) (Table 4). Icelinus burchami and I. tenuis also are found with I. borealis; however, the northernmost extent of their geographic ranges is Southeast Alaska and do not extend farther north into the Gulf of Alaska or into the Bering Sea (Matarese et al., 1989; Mecklenburg et al., 2002). Larvae of I. burchami and I. tenuis have not been identified, but there are subtle differences in meristic counts of juveniles and adults between these species and I. borealis (Table 4). Juvenile Icelinus may be distinguished by using adult characters in any geographic location (e.g., by the presence of elongated, threadlike first two dorsal spines in I. filamentosus). Icelinus quadriseriatus is the only species of Icelinus with currently identifiable and described early life history stages. Icelinus quadriseriatus is distributed from Sonoma County, California Sonoma County is on the northwest coast of California, one of the northernmost parts of the greater San Francisco Bay Area, U.S. Its population at the 2000 census was 458,614. Its largest city and county seat is Santa Rosa. , south to Cabo San Lucas Cabo San Lucas (popularly known as just Cabo) is a small city at the southern tip of the Baja California peninsula at , in the municipality of Los Cabos in the state of Baja California Sur, Mexico. , Baja California, Mexico (Feeney, 1987). Although I. borealis and I. quadriseriatus are geographically separated and their distributions do not overlap, it is important to compare the larvae of these species. Larvae of I. borealis and I. quadriseriatus are similarly pigmented; however they differ primarily in number of PVMs and ventral gut pigment. Icelinus borealis PVMs number from three to 14 (vs. 25-63). Icelinus borealis may have a few, individual melanophores present on the ventral gut during preflexion, whereas I. quadriseriatus has ventral gut pigment consisting of one to six rows of melanophores aligned anteroposteriorly in early development. Icelinus quadriseriatus retains ventral gut pigment throughout its larval development (Feeney, 1987). Icelinus borealis differs from I. quadriseriatus by having an analfin ray count of 11-14 (vs. 10-15), and a vertebral count of 35-36 (vs. 33-35) (Table 4). Icelinus borealis and I. quadriseriatus also undergo flexion at different times (8.0-11.0 mm vs. 5.2-7.6 mm, respectively) (Feeney, 1987). After examining all available putative larval specimens of Icelinus from the Bering Sea, it was found that the majority of larvae at AFSC were not I. borealis but probably members of the closely related genus, Icelus. The majority of larvae had higher myomere myomere /myo·mere/ (mi´o-mer) myotome (2). my·o·mere n. The segment within a metamere that develops into skeletal muscle. counts (37-42) than Icelus (35-36) and a different pelvic-fin count (1, 3) than I. borealis (1, 2) (Table 4). Larvae of I. borealis and Icelus had the same general body shape, presence of irregular PVMs (size, shape, location), similar pigmentation on the head, gut, and anus, four prominent preopercular spines, and a distinctive bony shelf on the anterior portion of the preopercle. Icelinus and Icelus were also placed in the same phenetic group by Richardson (1981) based on shared larval characters. There are five species of Icelus in the Bering Sea; however, Icelus spatula spatula /spat·u·la/ (spach´u-lah) [L.] 1. a wide, flat, blunt, usually flexible instrument of little thickness, used for spreading material on a smooth surface. 2. a spatulate structure. and I. spiniger are most abundant in the geographic area where Icelinus borealis is found (Matarese et al., 1989). This study provides a sound method for identifying larval I. borealis in the Bering Sea and is applicable to juvenile specimens as far south as southern Puget Sound, Washington. Although only two preflexion specimens were available for study, morphological characters and patterns of pigmentation at this stage of development are an important contribution. Taxonomic characters presented here could elucidate distinctiveness or similarity of Icelinus among other cottid genera (e.g., Ruscarius, Icelus) and co-occurring species (e.g., Icelinus filamentosus)--an important beginning to solving the complicated systematic relationships within the family Cottidae (Richardson, 1981). Although I. borealis larvae were identified in this study from the Bering Sea, definitive identification of larval I. borealis in other geographic areas will depend on the comparison of I. borealis with its congeners and other sympatric cottid larvae. Acknowledgments The author thanks J. Orr (AFSC), M. Busby, A. Matarese, and J. Napp for reviewing the manuscript. Data for adult groundfish and specimens of Icelinus and Icelus juveniles and adults were supplied by J. Orr and D. Stevenson (AFSC), who also granted use of the Resource Assessment and Conservation Engineering Groundfish Systematics Laboratory and digital radiograph radiograph /ra·dio·graph/ (-graf?) the film produced by radiography. ra·di·o·graph n. machine. J. Benson (AFSC) and M. Busby provided ArcView-GIS training and support. K. Maslenikov, University of Washington, provided radiographs and specimens of Icelus. This research is contribution EcoFOCI-0636 to the National Oceanic and Atmospheric Administration's Fisheries-Oceanography Coordinated Investigations. Manuscript submitted 18 September 2007. Manuscript accepted 29 October 2008. Fish. Bull. 107:175-185 (2009). The views and opinions expressed or implied in this article are those of the author and do not necessarily reflect the position of the National Marine Fisheries Service, NOAA NOAA abbr. National Oceanic and Atmospheric Administration Noun 1. NOAA - an agency in the Department of Commerce that maps the oceans and conserves their living resources; predicts changes to the earth's environment; . Literature cited Begle, D. P. 1989. Phylogenetic phy·lo·ge·net·ic adj. 1. Of or relating to phylogeny or phylogenetics. 2. Relating to or based on evolutionary development or history. analysis of the cottid genus Artedius (Teleostei: Scorpaeniformes). Copeia 1989:642-652. Blackburn, J. E. 1973. A survey of' the abundance, distribution and factors affecting distribution of ichthyoplankton in Skagit Bay. M.S. thesis, 136 p. Univ. Washington, Seattle, WA. Bolin, R. L. 1936. A revision of the genus Icelinus Jordan. Copeia 1936:151-159. Browne, P., J. L. Laake, and R. L. DeLong. 2002. Improving pinniped pinniped: see seal. pinniped Any member of the three existing families of aquatic, fin-footed mammals that constitute the suborder Pinnipedia (order Carnivora; see carnivore). diet analyses through identification of multiple skeletal structures in fecal samples. Fish. Bull. 100:423-433. Busby, M. S., and D. A. Ambrose. 1993. Development of larval and early juvenile pygmy poacher, Odontopyxis trispinosa, and blacktip poacher, Xeneretmus latifrons (Scorpaeniformes: Agonidae). Fish. Bull. 91:397-413. Feeney, R. F. 1987. Development of the eggs and larvae of the yellowchin sculpin, Icelinus quadriseriatus (Pisces: Cottidae). Fish. Bull. 85:201-212. Fritz, L. W., and S. Hinckley. 2005. A critical review of the regime shift--"junk food junk food n. Any of various prepackaged snack foods high in calories but low in nutritional value. junk food " --nutritional stress hypothesis for the decline of the western stock of Steller sea lion Noun 1. Steller sea lion - largest sea lion; of the northern Pacific Eumetopias jubatus, Steller's sea lion sea lion - any of several large eared seals of the northern Pacific related to fur seals but lacking their valuable coat . Mar. Mammal Sci. 21(3):476-518. Hoff, G. R. 2006. Biodiversity as an index of regime shift in the eastern Bering Sea. Fish. Bull. 104:226-237. Kendall, A. W., Jr., E. H. Ahlstrom, and H. G. Moser. 1984. Early life history of fishes and their characters. In Ontogeny ontogeny: see biogenetic law. Ontogeny The developmental history of an organism from its origin to maturity. It starts with fertilization and ends with the attainment of an adult state, usually expressed in terms of both maximal body and systematics of fishes, spec. publ. 1 (H. G. Moser, W. J. Richards, D. M. Cohen cohen or kohen (Hebrew: “priest”) Jewish priest descended from Zadok (a descendant of Aaron), priest at the First Temple of Jerusalem. The biblical priesthood was hereditary and male. , M. P. Fahay, A. W. Kendall Jr., and S. L. Richardson, eds.), p. 11-22. Am. Soc. Ichthyol. Herpetol., Allen Press, Lawrence, KS. Matarese, A. C., D. M. Blood, S. J. Picquelle, and J. L. Benson. 2003. Atlas of abundance and distribution patterns of ichthyoplankton from the Northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska Fisheries Science Center (1972-1996). U.S. Dep. Commer., NOAA Prof. Paper NMFS NMFS National Marine Fisheries Service NMFS National Mortality Followback Survey NMFS Network Multimedia File System NMFS Nested Mount File System 1, 281 p. Matarese, A. C., A. W. Kendall Jr., D. M. Blood, and B. M. Vinter. 1989. Laboratory guide to early life history stages of Northeast Pacific fishes. U.S. Dep. Commer., NOAA Tech. Rep. NMFS 80, 652 p. Matarese, A. C., and J. B. Marliave. 1982. Larval development of laboratory-reared rosylip sculpin, Ascelichthys rhodorus (Cottidae). Fish. Bull. 80:345-355. Mecklenburg, C. W., T. A. Mecklenburg, and L. K. Thorsteinson. 2002. Fishes of Alaska, 1037 p. Am. Fish. Soc., Bethesda, MD. Moser, H. G., R. L. Charter, P. E. Smith, D. A. Ambrose, S. R. Charter, C. A. Myer, E. M. Sandknop, and W. Watson. 1996. Distributional atlas of fish larvae and eggs in the California Current region. Calif. Coop. Oceanic Fish. Invest. Atlas 33. 1505 p. Scripps Inst. Ocean., La Jolla, CA. Mueter, F. J., and B. L. Norcross. 2000. Species composition and abundance of juvenile groundfishes around Steller sea lion Eumetopias jubatus rookeries in the Gulf of Alaska. Alaska Fish. Res. Bull. 7:33-43. Neira, F. J., A. G. Miskiewicz, and T. Trnski. 1998. Larvae of temperate Australian fishes--laboratory guide for larval fish identification, 474 p. Univ. Western Australia Press, Nedlands, Western Australia This article is about a suburb of Perth, Western Australia. For the local government area, see City of Nedlands. Nedlands is a western suburb of Perth, Western Australia. . Nelson, J. S. 2006. Fishes of the World Fishes of the World by Joseph S. Nelson is a standard reference for fish systematics. Now in its fourth edition, the work is a comprehensive overview of the 25,000-plus species known to science. , 4th ed., 601 p. John Wiley and Sons, Inc., Hoboken, NJ. Nelson, J. S., E. J. Crossman, H. Espinosa-Perez, L. T. Findley, C. R. Gilbert, R. N. Lea, and J. D. Williams
J.D. Williams (born May 22, 1978 in Newark, New Jersey) is an African-American actor notable for his starring roles in the HBO television programs Oz and The Wire. . 2004. Common and scientific names of fishes from the United States, Canada, and Mexico, 6th ed., 386 p. Am. Fish. Soc. Spec. Publ. 29, Bethesda, MD. Pietsch, T. W., and J. W. Orr. 2006. Triglops dorothy, a new species of sculpin (Teleostei: Scorpaeniformes: Cottidae) from the southern Sea of Okhotsk. Fish. Bull. 104:238-246. Potthoff, T. 1984. Clearing and staining techniques. In Ontogeny and systematics of fishes, spec. publ. 1 (H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall Jr., and S. L. Richardson, eds.), p. 35-37. Am. Soc. Ichthyol. Herpetol., Allen Press, Lawrence, KS. Richardson, S. L. 1977. Larval fishes in ocean waters off Yaquina Bay, Oregon: Abundance, distribution, and seasonality, January 1971-August 1972, 72 p. OSU (Open Source UNIX) Refers to the Unix variants that are maintained as open source, which were primarily BSD Unix and Linux until Sun made its Solaris operating system open source in 2005. Sea Grant Publ. ORESU-T-77-003. 1981. Current knowledge of larvae of sculpins (Pisces: Cottidae and allies) in Northeast Pacific genera with notes on intergeneric relationships. Fish. Bull. 79:103-121. Richardson, S. L., and W. G. Pearcy. 1977. Coastal and oceanic fish larvae in an area of upwelling off Yaquina Bay, Oregon. Fish. Bull. 75: 125-145. Richardson, S. L., and B. B. Washington. 1980. Guide to identification of some sculpin (Cottidae) larvae from marine and brackish brack·ish adj. 1. Having a somewhat salty taste, especially from containing a mixture of seawater and fresh water: "You could cut the brackish winds with a knife/Here in Nantucket" waters off Oregon and adjacent areas in the Northeast Pacific. NOAA Tech. Rep. NMFS 430, 56 p. [With errata er·ra·ta n. Plural of erratum. sheet dated May 1981.] Rosenblatt, R. H., and W. L. Smith. 2004. Icelinus limbaughi: A new species of sculpin (Teleostei: Cettidae) from southern California. Copeia 2004:556-561. Tsuruoka, O., T. Abe, H. Munehara, and M. Yabe. 2006. Record of a cottid fish, Icelinus pietschi, collected from Hokkaido and Miyagi Prefecture, Japan. Jap. J. Ichthyol. 53(1):89-93. Washington, B. B. 1981. Identification and systematics of larvae of Artedius, Clinocottus, and Oligocottus (Scorpaeniformes: Cottidae). M.S. thesis, 202 p. Oregon State Univ., Corvallis, OR. Yabe, M., S. Maruyama, and K. Amaoka. 1983. First records of five cottid fishes and a psychrolutid fish from Japan. Jap. J. Ichthyol. 29(4):456-464. Yabe, M., A. Soma soma (sō`mə), psychotropic plant, the juice of which was sometimes drunk as part of the Vedic sacrifice (see Veda). Many hymns in the Rig-Veda are in praise of soma. , and K. Amaoka. 2001. Icelinus pietschi sp. nov. and a rare species, Sigmistes smithi, from the southern Kuril Archipelago (Scorpaeniformes: Cottidae). Ichthyol. Res. 48(1):6570. Yabe, M., K. Tsumura, and M. Katayama. 1980. Description of a new cottid fish, Icelinus japonicus, from Japanese waters. Jap. J. Ichthyol. 27(2):106-110. Rachael L. Cartwright Email address: rachcart@hotmail.com National Marine Fisheries Service, NOAA Alaska Fisheries Science Center 7600 Sand Point Way NE, Building 4 Seattle, Washington 98115
Table 1
Body proportions of larvae and juveniles of northern sculpin
(Icelinus borealis). Values for each body proportion are
expressed as percentage of standard length (SL) or head
length (HL): mean, standard deviation, and range.
Body proportion Flexion
Sample size 9
Standard length 9.1 [+ or -] 0.74 (8.0-10.2)
Head length/SL 25.2 [+ or -] 0.02 (22.2-28.4)
Snout length/HL 27.9 [+ or -] 0.04 (24.8-37.9)
Eye diameter/HL 31.0 [+ or -] 0.03 (25.3-33.7)
Snout-to-anus length/SL 44.6 [+ or -] 0.03 (38.7-48.6)
Body depth/SL 20.3 [+ or -] 10.02 (17.6-23.2)
Pectoral-fin length/SL 10.1 [+ or -] 0.02 (6.6-13.4)
Body proportion Postflexion
Sample size 27
Standard length 14.9 [+ or -] 12.40 (11.1-22.7)
Head length/SL 34.3 [+ or -] 0.04 (26.2-41.5)
Snout length/HL 26.8 [+ or -] 0.03 (20.0-32.4)
Eye diameter/HL 24.3 [+ or -] 0.02 (20.2-30.3)
Snout-to-anus length/SL 47.2 [+ or -] 0.03 (42.3-54.0)
Body depth/SL 21.9 [+ or -] 0.02 (17.6-27.1)
Pectoral-fin length/SL 24.2 [+ or -] 4.30 (13.8-32.4)
Body proportion Juvenile
Sample size 8
Standard length 38.6 [+ or -] 10.3 (24.1-51.7)
Head length/SL 37.9 [+ or -] 0.02 (35.2-39.8)
Snout length/HL 27.9 [+ or -] 0.05 (21.8-39.0)
Eye diameter/HL 27.8 [+ or -] 0.01 (26.1-30.6)
Snout-to-anus length/SL 50.6 [+ or -] 0.04 (46.4-57.7)
Body depth/SL 20.5 [+ or -] 0.02 (17.4-22.7)
Pectoral-fin length/SL 24.5 [+ or -] 12.80 (19.6-28.8)
Table 2
Total postanal ventral melanophores (PVMs) of larvae
and juveniles of northern sculpin (Icelinus borealis).
Specimens between dotted lines (----) are undergoing
notochord flexion; specimens between lines (---) are
in transformation stage. Abbreviation: SL = standard
length.
Body Postanal
length ventral
(mm SL) melanophores
7.4 --
7.9 9
8.8# 9#
10.2# 4#
11.6 9
13.4 14
14.3 9
14.8 9
14.9 7
15.1 11
15.8 9
16.0@ 8@
16.3@ 10@
17.9@ 7@
19.6@ 5@
22.7@ 3@
24.1 3
24.9 7
32.1 0
41.7 0
43.2 0
45.1 0
45.9 0
51.7 0
Note: Undergoing notochord flexion
indicated with #.
Note: Transformation stage indicated with @.
Table 3
Meristic counts of larvae and juveniles of northern sculpin (Icelinus
borealis). Specimens between dotted lines (-----) are undergoing
notochord flexion; specimens between dashed lines (----) are
undergoing transformation. Abbreviation: SL = standard length.
Body Dorsal-fin
length Anal-fin
(mm SL) Spines Rays rays
7.4 -- -- --
7.9 -- -- --
* 8.8 (a) --(a) --(a) --(a)
* 10.2 (a) --(a) --(a) --(a)
* 11.6 -- -- --
* 13.4 -- -- --
* 14.3 -- -- 13
* 14.8 -- -- --
* 14.9 -- -- --
* 15.1 -- -- --
* 15.8 X 17 14
* 16.0 (b) -- (b) -- (b) -- (b)
* 16.3 (b) X (b) 16 (b) 13 (b)
+ 17.9 (b) X (b) 16 (b) # (b)
* 19.6 (b) XI (b) 16 (b) 12 (b)
* 22.7 (b) XI (b) 16 (b) 13 (b)
* 24.1 X 16 13
+24.9 IX 16 13
+32.1 X 16 13
+41.7 IX 16 13
+43.2 X 16 13
+45.1 X 15 12
+45.9 # 15 12
+51.7 IX 14 12
Pectoral-fin Pelvic-fin
Body spine & rays spine & rays
length
(mm SL) Left Right Left Right
7.4 -- -- -- --
7.9 -- -- -- --
* 8.8 (a) -- -- -- --
* 10.2 (a) -- -- -- --
* 11.6 -- -- -- --
* 13.4 -- -- -- --
* 14.3 16 16 I, 2 I, 2
* 14.8 -- -- -- --
* 14.9 16 16 -- --
* 15.1 16 16 -- --
* 15.8 16 17 I, 2 I, 2
* 16.0 (b) 16 (b) 16 (b) I, 2 (b) I, 2 (b)
* 16.3 (b) 16 (b) 16 (b) I, 2 (b) I, 2 (b)
+ 17.9 (b) 15 (b) 15 (b) I, 2 (b) I, 2 (b)
* 19.6 (b) 16 (b) 16 (b) I, 2 (b) I, 2 (b)
* 22.7 (b) 16 (b) 15 (b) I, 2 (b) I, 2 (b)
* 24.1 16 16 I, 2 I, 2
+24.9 16 16 I, 2 I, 2
+32.1 16 16 I, 2 I, 2
+41.7 16 16 I, 2 I, 2
+43.2 16 16 I, 2 I, 2
+45.1 16 16 I, 2 I, 2
+45.9 15 15 l, 2 I, 2
+51.7 16 16 I, 2 I, 2
Caudal-fin rays
Body Superior Inferior
length
(mm SL) Procurrent Principal Procurrent Principal
7.4 -- -- -- --
7.9 -- -- -- --
* 8.8 (a) -- -- -- --
* 10.2 (a) -- -- -- --
* 11.6 -- -- -- --
* 13.4 -- -- -- --
* 14.3 -- 6 6 7
* 14.8 -- 6 6 --
* 14.9 -- -- -- --
* 15.1 -- -- -- --
* 15.8 9 6 6 7
* 16.0 (b) -- (b) 6 (b) 6 (b) -- (b)
* 16.3 (b) 10 (b) 6 (b) 6 (b) 8 (b)
+ 17.9 (b) 11 (b) 6 (b) 6 (b) 8 (b)
* 19.6 (b) 10 (b) 6 (b) 6 (b) 7 (b)
* 22.7 (b) 10 (b) 6 (b) 6 (b) 4 (b)
* 24.1 10 6 6 8
+24.9 9 6 6 8
+32.1 10 6 6 7
+41.7 10 6 6 7
+43.2 8 6 6 6
+45.1 9 6 6 8
+45.9 9 6 6 8
+51.7 9 6 6 8
Caudal-fin rays
Vertebrae
Body Branchio-
length stegal
(mm SL) Abdominal Caudal Total rays
7.4 -- -- -- --
7.9 -- -- -- --
* 8.8 (a) -- -- -- --
* 10.2 (a) -- -- -- --
* 11.6 -- -- -- --
* 13.4 -- -- -- --
* 14.3 9 26 35 6
* 14.8 -- -- -- --
* 14.9 -- -- -- 6
* 15.1 -- -- -- 6
* 15.8 10 25 35 6
* 16.0 (b) -- (b) -- (b) -- (b) 6 (b)
* 16.3 (b) 11 (b) 25 (b) 36 (b) 6 (b)
+ 17.9 (b) 11 (b) 25 (b) 36 (b) 6 (b)
* 19.6 (b) 10 (b) 25 (b) 35 (b) 6 (b)
* 22.7 (b) 10 (b) 25 (b) 35 (b) 6 (b)
* 24.1 # # 36 6
+24.9 10 26 36 6
+32.1 10 26 36 6
+41.7 11 25 36 6
+43.2 11 25 36 6
+45.1 # # 36 6
+45.9 11 25 36 6
+51.7 11 25 36 6
* Cleared and stained specimens.
+ Radiographed specimens.
# Could not obtain accurate count.
Table 4
Comparison of meristic counts of Icelinus and Icelus
(Yabe et al., 1980; Matarese et al., 1989; Moser et
al., 1996; Yabe et al., 2001; Mecklenburg et al.,
2002; Rosenblatt and Smith, 2004; Tsuruoka et al.,
2006). Counts in parentheses (I. borealis) indicate
the mode. Abbreviations: C=central; S=southern;
SE=southeast; SS=south of southern.
Species Common name Distribution
Icelinus borealis northern sculpin Washington-Bering Sea
I. burchami dusky sculpin S California-SE Alaska
I. cavifrons pit-head sculpin SS California-
C California
I. filamentosus threadfin sculpin S California-
Gulf of Alaska
I. fimbriatus fringed sculpin S California-
British Columbia
I. japonicas Futasuji-kajika Japan
I. Limbaughi canyon sculpin S California
I. oculatus frogmouth sculpin S California-
British Columbia
I. pietschi Hime-futasuji-kajika Onagawa, Japan-
Iturup L, Kuril Is.
I. quadriseriatus yellowchin sculpin SS California-
C California
I. tenais spotfin sculpin SS California-
SE Alaska
Icelus canaliculatus blacknose sculpin Gulf of Alaska-
Bering Sea
I. earyops wide-eye sculpin Gulf of Alaska-
Bering Sea
1. spatula spatulate sculpin Gulf of
Alaska-Arctic
I. spiniger thorny sculpin British Columbia-
Bering Sea
I. uncinalis uncinate sculpin Bering Sea
Fins
Species Dorsal Anal
Icelinus borealis IX-XI + 14-17 (X + 16) 11-14 (13)
I. burchami VIII-XI + 15-18 10-14
I. cavifrons IX-XII + 12-15 11-13
I. filamentosus IX-XII + 15-18 13-16
I. fimbriatus X-XI + 12-14 12-14
I. japonicas IX-X + 12-13 10-11
I. Limbaughi IX-X + 13-15 8-12
I. oculatus X-XI + 15-17 13-14
I. pietschi X + 13-14 11-12
I. quadriseriatus VII-X + 12-16 10-15
I. tenais IX-XI + 16-19 13-17
Icelus canaliculatus VII VIII + 22-25 18-20
I. earyops VIII-X + 20-23 15-19
1. spatula VII-XI + 18-22 13-18
I. spiniger VIII-X + 19-23 15-19
I. uncinalis IX + 19-20 14-16
Fins
Species Pectoral Pelvic Vertebrae
Icelinus borealis 14-17 (16) I,2 35-36 *
I. burchami 16-19 I,2 33-37
I. cavifrons 14-16 I,2 35-37
I. filamentosus 16-18 I,2 34-37
I. fimbriatus 16-18 I,2 35-37
I. japonicas 15-17 I,2 33
I. Limbaughi 15-17 I,2 31-36
I. oculatus 17 I,2 37
I. pietschi 16 I,2 32-34
I. quadriseriatus 15-17 I,2 33-35
I. tenais 15-17 I,2 37-39
Icelus canaliculatus 15-19 I,3 37-39
I. earyops 16-18 I,3 41-42 **
1. spatula 16-20 I,3 39-41
I. spiniger 17-20 I,3 40-42
I. uncinalis 17-18 I,3 37-40
* Count, obtained from literature and radiographs of adult specimens.
** Counts obtained from radiographs of adult specimens.
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