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Matarese.Pdf LARVAL DEVELOPMENT OF PACIFIC TOMCOD, MICROGADUS PROXIMUS, IN THE NORTHEAST PACIFIC OCEAN WITH COMPARATIVE NOTES ON LARVAE OF WALLEYE POLLOCK, THERAGRA CHALCOGRAMMA, AND PACIFIC COD, GADUS MACROCEPHALUS (GADIDAE) l ANN C. MATARESE,' SALLY L. RICHARDSON: AND JEAN R. DUNN ABSTRACf A developmental series from yolk-sac larvae through juveniles (2.7-46.6 mm SL) of Microgadus proximus from the northeast Pacific Ocean is described and illustrated. Larvae hatch at approximately 2.7 mm SL and the yolk is absorbed by 3.0 mm S1. Notochord flexion begins between 8.0 and 10.0 mm SL, is completed at 15.0 mm SL, and transformation occurs between 22.0 and 28.0 mm S1. Larvae have specific pigment patterns, particularly in the postanal region where melanophores are arranged in two bars, anterior and posterior. At 5.0-6.0 mm 8L, the anterior bar is located at 41-53% SL and the posterior bar is at 61-74% SL. Melanophore patterns on the head, gut, and caudal region also distin­ guish M. proximus. The occurrence oflarvae taken within 18 km ofshore off Oregon during plankton surveys in 1971-72 is discusssed, and data indicate a winter-spring spawning period. A combination of pigmentation characters, primarily the length and position of the anterior and posterior pigment bars, and meristic counts can distinguish larvae of M. proximus from Theragra chalcogramma and Gadus macrocephalus. The anterior bar is located at 47-55% SL in T. chalco­ gramma and at 40-57% SL in G. macrocephalus. The posterior bar is located at 69-79% SL in T. chalcogramma and at 59-81% SL in G. macrocephalus. Both T. chalcogramma and G. macrocephalus have 4 rays on the superior hypural element while M. proximus has 5 rays. Other characters useful in separating the three species include head, gut, mediolateral, postanal, and caudal pigment and stripe continuity. Problems have been encountered in distinguish­ larvae of these three gadids in mixed samples. inglarvaeofPacific tomcod, Microgadus proximus Eggs and larvae of the Atlantic tomcod, Micro­ (Girard); walleye pollock, Theragra chalco­ gadus tomcod, were described by Booth (1967), but gramma (Pallas); and Pacific cod, Gadus mac­ eggs of M. proximus are unknown. rocephalus (Tilesius), in samples from the north­ The geographic range of M. proximus extends eastern Pacific Ocean where these three species from off central California (Isaacson 1965) to the might cooccur (Waldron 1972; Dunn and Naplin3). Gulf ofAlaska and Unalaska Island (Wilimovsky Larvae of T. chalcogramma were described by 1964). Their presence in the Bering Sea remains Gorbunova (1954) and Yusa (1954). Larvae of G. unconfirmed, although they were reportedtooccur (morhua) macrocephalus larvae were described by in the Bering Sea by Tanner (1894) and Hart Gorbunova (1954), Uchida et al. (1958), and (1973). Microgadus proximus was not captured in Mukhacheva and Zviagina (1960). In this report the eastern Bering Seaduring extensive multives­ 4 we provide the first published description of the sel groundfish surveys in 1974 (Pereyra et a1. ) or 5 larvae of M. proximus and comparative material in 1976 (Bakkala and Smith ). Microgadus prox­ on larvae of T. chalcogramma and G. mac­ imus is found from near-surface waters to ap- rocephalus that should enable workers to identify 'Pereyra, W. T., J. E. Reeves, and R. G. Bakkala. 1976. De­ INorthwest and Alaska Fisheries Center, National Marine mersal fish and shellfish resources of the eastern Bering Sea in Fisheries Service, NOAA, 2725 Montlake Boulevard East, the baseline year 1975. Unpubl. manuscr., Vol. 1,619 p.; Vol. Seattle, WA 98112. 2, 534 p. Northwest and Alaska Fisheries Center National 2Gulf Coast Research Laboratory, East Beach Drive, Ocean Marine Fisheries Service, NOAA, 2725 Montlake' Blvd. E., Springs, MS 39564. Seattle, WA 98112. 3Dunn, J. R., and N. A. Naplin. 1974. Fish eggs and larvae sBakkala, R. G., and G. B. Smith. 1978. Demersal fish re­ collected from waters adjacent to Kodiak Island, Alaska, during sources of the eastern Bering Sea: Spring 1976. Unpub!. April and May, 1972. Unpubl. manuscr., 61 p. Northwest and manuscr., Vol. 1, 234 p.; Vol. 2, 404 p. Northwest and Alaska Alaska Fisheries Center, National Marine Fisheries Service, Fisheries Center, National Marine Fisheries Service, NOAA, NOAA, 2725 Montlake Blvd. E., Seattle, WA 98112. 2725 Montlake Blvd. E., Seattle, WA 98112. Manuscript accepted May 1980. FISHERY BULLETIN: VOL. 78, NO.4, 1981. 923 FISHERY BULLETIN: VOL. 78. NO.4 proximately 220 m (Miller and Lea 1972). adult T. chalcogramma specimens at NWAFC and Planktonic larvae of M. proximus were the from the Institute of Animal Resource Ecology, dominant gadid and fourth most abundant taxon University of British Columbia, Vancouver. in a coastal assemblage offish larvae occurring off Illustrations oflarvae were made with the aid of Yaquina Bay, Oreg., in 1971-72 (Richardson and a camera lucida. All specimens had been preserved Pearcy 1977). in either 3-5% Formalin,6 buffered with sodium Information on the biology and taxonomy of M. borate, or 95% ethanol. Illustrations of caudal fin proximus is limited. Schultz and Welander (1935) development were drawn from cleared and stained presented counts of meristic structures and mor­ specimens. phological measurements of M. proximus. Svetovidov (1948), in his review of Gadiformes, Measurements provided some meristic data and a brief os­ teological description ofM. proximus based on two The following measurements were made on 72 skeletons. Briefnotes on the biology ofthe species unstained larvae and juveniles (2.7-46.6 mm SL) are included in Clemens and Wilby (1961), Miller of M. proximus using an ocular micrometer in a and Lea (1972), and Hart (1973). It is not of com­ stereomicroscope: mercial importance in part because of its small size (to 30 em) but it is taken in recreational Standard length (SL)-Snout tip to notochord tip catches (Beardsley and Bond 1970; Hart 1973). prior to development of caudal fin, then to pos­ terior margin of hypural element. (All body METHODS lengths in this study are standard lengths.) Head length (HL)-Snout tip to posterior edge of Specimens opercle (to pectoral fin base in yolk sac and very small larvae before opercular margin is visible). Several hundred larvae ofthe three species were Snout length-Snout tip to anterior margin of examined in the course of this study. Microgadus orbit of left eye. proximus larvae andjuveniles were obtained from Upper jaw length-Snout tip to posterior margin plankton and trawl samples collected off Oregon of maxillary. in 1971-1973 and 1979, by the School ofOceanog­ Eye diameter-Greatest diameter of left orbit. raphy, Oregon State University (OSU), Corvallis, Body depth at pectoral-Vertical distance from and off Washington in 1972 by the Northwest and dorsal to ventral body margin at pectoral fin Alaska Fisheries Center (NWAFC). Additional base. specimens were obtained from plankton collected Body depth at anus-Vertical distance from dor­ inPugetSound, Wash., in 1977-79 bythe Fisheries sal to ventral body surface at center of anal Research Institute (FRI), University of Washing­ opening. ton, Seattle; and in 1978 by Ecology Consultants, Snout to anus-Distance along body midline from Inc., Fort Collins, Colo. Larvae were also collected snout tip to vertical through center ofanal open­ from near Kodiak Island, Alaska, in 1977-79 by ing. FRI. Snout to first dorsal fin-Distance along the body Theragra chalcogramma larvae were collected midline to vertical through origin of anterior­ in the eastern Bering Sea (1971,1976-78) and off most dorsal fin ray of first dorsal fin. Kodiak Island (1972, 1977-78) by NWAFC. Ad­ Snout to second dorsal fin-Distance along body ditional specimens from Puget Sound, Wash., midline to vertical through origin of anterior­ (1977,1978) were provided by FRI and by Ecology most fin ray ofsecond dorsal fin. Consultants, Inc., and from off Kodiak Island Snout to third dorsal fin-Distance along body (1978) by FRI. midline to vertical through origin of anterior­ Gadus macrocephalus larvae were collected most fin ray ofthird dorsal fin. near Kodiak Island in 1978 by NWAFC and FRI Snout to first anal fin-Distance along body mid­ and from Puget Sound, Wash., in 1977-79 by FRI. line to vertical through origin of anteriormost Radiographs were examined of juvenile and fin ray of first anal fin. adult M. proximus and G. macrocephalus speci­ mens in the collections in the Department of "References to trade names do not imply endorsement by the Fisheries and Wildlife, OSU, and NWAFC, and of National Marine Fisheries Service, NOAA. 924 MATARESE ET AL.: LARVAL DEVELOPMENT OF PACIFIC TOMCOD Snout to second anal fin-Distance along body development of dorsal and ventral procurrent midline to vertical through origin of anterior­ caudal rays before the development of hypurals. most fin ray ofsecond anal fin. Considering the collection locations (coastal Oregon and Washington) and the range of myo­ Meristic Structures mere counts, the specimens could have been one of only three potential species, M. proximus, T. chal­ Counts of meristic structures were made on 128 cogramma, or G. macrocephalus. Meristic charac­ stained M. proximus larvae and juveniles. Sixty­ ters in the literature (e.g., Svetovidov 1948; Miller six specimens (3.1-41.1 mm SL) were cleared and and Lea 1972; Hart 1973) are inadequate to sepa­ stained with Alizarin Red S (Taylor 1967), and 62 rate all three species. Additional counts obtained larvae (5.1-23.4 mm SL) were stained with Aliza­ in this study, particularly caudal vertebrae and fin rin Red and Alcian Blue (Dingerkus and Uhler rays on the superior hypural (Tables 1, 2), enabled 1977). Both series ofstained samples were used to positive identification of the series as M. prox­ make counts of meristic structures and to deter­ imus. mine the onset and sequence ofossification as in­ dicated by the uptake ofAlizarin Red.
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