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Species Diversity, 2007, 12, 17-27

Pelagic Juveniles of Two Morids (Teleostei: Gadiformes:

Moridae), Antimora microlepis and Ph{ysiculusjaponicus,

from the Western North Pacific

Makoto Okamotoi, Naoshi Sato2, TakashiAsahida2 and Yoshiro Watanabe3

' 7bhoku IVlritional Fisheries Research institute, 3-275 Shinhama-cho, Shiogama, Mlyagi, g85-OOOI .Jtipan E-nzaig: okamaho@,cij[rrc.go.jp 2Schoot ofFisheries Sciences, Kitasato U}iiversity, 1604 Okirai, Sanriku-cho, Qfttnato, Iteate, a22-OIOI .1lrzpan 3Ocean Research Iitstitute, Lbeiversily of 7bdyo, Minamida4 IVtikano-ku, 7bdyo, 164-8639 Jopan

(Received 1 July 2006: Accepted 20 November 2006)

Pelagic juveniles of two morids, Antimora microlepis Bean, 1890 and Pbysiculus j(rponicus Hilgendort 1879, were colleeted by midwater t.rawl (O-20m depth) frem transition waters between the Oyashio and Kuroshio fironts ef the western North Pacific in May, 1989. Juveniles of A. microtqpis (13 specimens. 30.2-54,3mm standard length, SL> characteristieally have an elengated body, posteriorly, positioned anus, 24-25 precaudal vertebrae, chin with a barbel, no ventral luminous organ, elongated pelvic fin rays, and a non-protruding snout. Juveniles of P. jmponicus (six specimens, 20.4-39.0mm SL) characteristically have an elongated body, chin with a barbel, ventral lu- minous organ anterior to the anus, elongated pelvic fin rays, a pointed cau- dal fin, glO dorsal fin rays, 6070 second dorsal fin rays, 6&72 anal fin rays, six pelvic fin rays, 16=41A2 vertebrae, and ca. 130 lengitudinal scales. Key Words: Teleostei, Gadiformes, Moridae, Antimora microlepis, Pdysicu- lus j'mponicus, pelagic juvenile,

Introduction

The family Moridae currently comprises 18 genera and about 110 specjes, which occur widely from coastal to pelagic waters over the entire world except in the Arctic Ocean (Paulin 1989a; Okamura 1995; Eschmeyer 1998). The adult mor- phology and taxonomy of the family have been comprehensively studied by Paulin (1983, 1989a) and Cohen (1984), The family is defined by the fo11owing four charac- ters: a swim bladder-auditory capsule connection, a caudal skeleton with fbur or five hypurals and X-Y bones, a joined first neural spine, and distinctive otoliths, Occurrence and merphological infoumation on the early life stages of the family are limited to about 20 species in nine genera (Eretmqphorus, Gadelta, Guttigadus, Halangyreus, Laemonema, Mora, Pdysiculus, Rhynchogadus, and Svetovidovia) (Fahay and Markle 1984; Cohen 1986; Okiyama 1988; Markle 1989; Olivar and Fortufio 1991; Ambrose 1996; Melendez and Markle 1997; Fukui et al. 2003; Comyns

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140eE 14se 15oe asse 45[N

4oe

35e

Fig. 1. Map showing sampling sites of pelagic juveniles of Antimora microlepis (circle) and Pdysiculttsjaponicus(triangle),

and Bond 2005). Researchers at the Tohoku National Fisheries Research Institute have been studying the larvae and juveniles of fish species that serve as fisheries resources in the transition waters between the Oyashio and Kuroshio fronts of the western North Pacific, During a cruise to the transition waters, pelagie juvenile specimens of morids were collected by midwater trawl at twe sampling stations on 16 and 18 May 1989 (Fig. 1). We identMed two species from these juvenile samples, Antimora microlepis Bean, 1890 and Pdysiculus joponicus Hilgendorf, 1879. In this paper, the pelagic juveniles ofA, microlqpis and P. japonicus are described fbr the first time.

Materials and Methods

Thirteen juvenile specimens (30.2-54.3mm SL) of Antimora microlepis were collected at 39eOO'N, 149000'E in the transition waters off Iwate Prefecture on 16 May 1989 (Fig. 1). Six juvenile speeimens (20.4-39.0mm SL) of PIrysiculus jcrponic"s were collected from 38000'N, 151UOO'E in the transition waters off Miyagi Prefeeture on 18 May 1989 (Fig. 1). At both sampling stations midwater trawls were towed at approximate]y 2 knots in the O-20m ]ayer. A conductivity, temperature, and depth prefiler (CTD) cast was made at all sampling stations down to 1000m depth. The Oyashio and Kuroshio fronts were defined by the 50 and 150C isotherms at 100m depth, respectively, and the transition waters were those located between the fronts (Kawai 1972; Odate 1994). Temperatures in the O-20m layer at the two sam-

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Pelagic juveniles of two Pacific morids 19

pling stations off Iwate and Miyagi Prefectures were 11.4-13,4"C and 11,9-14,50C, re- spectively. The pelagic juvenile specimens were fixed in 10% buffered fbrmalin and Iater transferred to 70e,6 ethanol. Measurement methods, abbreviations, and definitiens fbllew Pauljn (1983). For observations of skeletal features, three speci- mens of A, microlqpis (42.6-50.9mm SL) and two specimens of P. jqponicus (25.1-30,2mm SL) were cleared and stained fo11owing Potthoff (1984). All the speci- mens examined in this study are deposited in the School of Fisheries Sciences, Ki- tasato University (FSKU).

Taxonomic Account

Antimora microlepis Bean, 1890 (Fig. 2)

Material examined. Pelagic juvenjles: 13 specimens, FSKU-P 19841-19853, 30,2-54.3mm SL, off Iwate Prefecture, northeastern Japan, 39000'N, 149000'E, O-20

m, 16 May 1,989. Identification. Thirteen juvenile specimens were assigned to the genus An- timora of the Moridae because they had all of the following characters: posteriorly positioned anus, 24-25 precaudal vertebrae, chin with a barbel, and no ventral lu- minous organ (Paulin 1983; Cohen 1984). Aduit specimens of Anttmora species are easily distinguished from those of other genera of morids by their protruding snout, but the snout of pelagic juveniles is rounded (see Remarks). The genus An- timora is represented in the North Pacific region by only A. microlepis (cf, Small 1981). Description. Meristic counts and measurements given in Table 1. Body elon- gate, compressed, gradually tapering to caudal peduncle, greatest depth at pectoral fin base. Head rounded, dorsum slightly elevated. Mouth moderate in size, not pro- truded; maxilla reaching anterior margin of eye. One or two rows of small conical teeth on premaxil!a and dentary; no teeth on prevomer and palatines, Snout rounded with two nostrils, each nearly equal to eye diameter. Eye of moderate size, dorsally situated on head with dorsal margin close to dorsal profile. Interorbital space elevated, width longer than eye diameter. Short barbel present on chin. Gil1 opening wide, opercular region thin, No ventral luminous organ, Anus located pos- teriorly, even with 16th-18th rays of second dorsal fin. First dorsal fin origin

'

Fig. 2. Pelagic juvenile ofAntimora microlepis, 49,5mm SL, FSKU-P 19842. Scale bar: 5mm.

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Table 1. Meristic counts and proportional measurements of juveniles of Antimora nzicrolepts and Plavsiculus jmponicus.

Species (number of specimens) Antimora microlepis (13) Physiculusjcrponictts (6) Standard length (mm) 30,2-54.3 20.4-39.0

Counts

First dorsal fin rays 4-5 9-1066-7068-7224-25 Second dorsal fin rays 5C52 Anal fin rays 38-40 Pectora] fin rays 18-19 Pelvic fin ravs 6 6 Branchiostegal rays 74-5-1 73+7-8 Gill rakers 10-12 Precaudal vertebrae 2425 164142 Caudal vertebrae 3436

ftt"ttkt"tt ± t ±t ±± ±t ± ±tde" ± "kttkt""++dede"tde""t"de""dedede"dededet"+-+

Measurements (V) of standard length) Head Iength 23,1- 25.5 23.0-24.6 Body depth at pectoral fin 20,6-23.6 21.4-23.3 Bocty depth at anus 12.1-15.9 22.1-25,9 Predorsal length 26.7-28.8 30.9-31,8 Preanus length 52.]-55.6 23.5-25.9 Pectoral fin length 24.0-26.3 18.5-21,1 Pelvic fin length 19.3-25.1 24.2-27.5 First dorsal fin base 3,3-4.8 8.0-9.660.7-62.8 Seeond dorsal fin base 61,5-66.9 Ana] fin base 35.7-36.8 52.0-58.9 Caudal peduncle depth 3,or3.6 2.6-3.2 Measurements (V: ofhead ]ength) Eye diameter 28.1-33,2 32.2-39,O Snout length 24.1- 26.3 16.4-21.1 Upper jaw length 37.3-44.2 40.0-44.4 Barbel length 4.2-6,3 8.2-11,2

above posterior margin of opercle; no elongated rays. Second dorsal fin origin just behind first dorsal fin; ]ast ray not connected to caudal fin by membrane. Anal fin origin a short distance behind anus, approximately below midlength peint of sec- ond dorsal fin base, Pectoral fin large, inserted below first dorsal fin origin. Pelvic fin with flat base, 2nd to 6th rays filamentously elongated, Caudal fin pointed, free from dorsal and anal fins, Three shallow Iongitudinal lines on lateral side of body. Scales undeveloped. Pignzentation in alcohol. Body color mainly light brown, with many large and small melanophores scattered on surface of body except caudal peduncle region. Large melanopheres in opercular region and on base of pectoral fin, Surface of brain case and snout, internal peritoneum, and dorsal and anal fin membranes covered with smal1 melanophores. Pectoral and caudal fins unpigmented. Remarks. The fin ray counts and the posterior position of the anus in the present juvenile specimens agree with two species of morids inhabiting the North Pacific, Antimora microlepis and Lapidion schmidti Svetovidov, 1936, but the pres- ent juveniles have 24-25 preeaudal vertebrae (based on cleared-and-stained speci-

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Pelagic juveniles of two Pacific merids 21

mens), a unique character of Antinzora among the morid genera (Fahay and Markle 1984; Paulin 1989a, 1989b; Paulin et aL 1989). The number of precaudal ver- tebrae in L. schrnidti i's 17-18 (Nakaya et al, 1980) many fewer than in Antimora. Al- though juveniles of L, schmidti have been collected in New Zealand (Paulin 1989a), no juveniles of any species of Lqpidion have yet been described and illustrated (Templeman 1970; Comyns and Bond 2005), Although adults of Antimora microlepis have a protruding snout tip, the sneut

ofjuveniles is rounded, Morphological difference in this character between adult and juvenile is well known in the Macrouridae, In this family the snout tip, fins and pigmentation change rapidly during metamerphosis (Merrett 1989; Olivar and Fortufio 1991; Ambrose 1996). Speeimens of A. microlepis smaller than 100mm SL are rare (Iwamoto 1975; Small 1981). We confirrned that a specimen of 118mm SL (FSKU-P 19677) has a protruded snout tip, and the largest juvenile specimen we ex- amined is 54.3mm SL; therefore, we conclude that the ontogenetic change in the

shape of the snout tip in A. nzicrolopis occurs between ca. 60 and 100mm SL, In some speeies of the morid genera Gadella and Laemonema the number of pelvie fin rays becomes reduced during ear]y life stages (Ambrose 1996; MelOndez and Markle 1997; Comyns and Bond 2005), In contrast, in A, microlqpis the number of pelvic fin rays in juveniles and adults is the same, There are two valid species of the genus Antimora, i.e., A. microlepis and A. rostrata (GUnther, 1878); in both species, the eggs, larvae, and juveniles have all been unknown until now (Ambrose 1996; Comyns and Bond 2005). This study is the first report of early life stages in this genus. Although sexual maturity and the dis- tribution pattern of adult A. microlepis have been poorly studied, Wenner and Mu- sick (1977) supposed that A. rostrata inhabiting the western North Atlantic may migrate to and spawn in waters deeper than 3000m in the northern part of the area.

Ptrysiculusjmponicus Hilgendorf, 1879 (Fig. 3)

Material examined. Pelagic juveniles: 6 specimens, FSKU-P 19854-19859, 20.4-39.0mm SL, eff Miyagi Prefecture, northeastern Japan, 38000'N, 151oOO'E, O-20

m, 18 May 1989, Identification. Six juvenile specimens were assigned to the genus PhysicuZus because they had all of the fo11owing characters: a compressed and elongated body,

two dorsal fins and one anal fin, a separate caudal fin, the chin with a barbel, a ven- tral luminous organ anterior to the anus (Fig. 3B), and no vomerine teeth (Paulin 1983, 1989b). The genus Physicultts in the western North Pacific region includes six species (Paulin 1989b)i P. chigodarana Paulin, 1989, P. aynodon Sazonov, 1986, P. jqponicus, P, nigripinnis Okamura, 1982, P. rhodQpinnis Okamura, 1982, and P. yoshidae Okamura, 1982, The present juvenile specimens have 9-10 first dorsal fin rays, 66-70 second dorsal fin rays, 68-72 anal fin rays, six pelvic fin rays, 16+4142 vertebrae, and ca. 130 Iongitudinal scales, These values agree only with those of P, jmponicus among the six western North Pacific species of PItysicutus (Table 2). Description. Meristic counts and measurements given in Table 1. Body elon- gated, compressed, tapering to very nanrow caudal peduncle, greatest depth at

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B

-

Fig. 3. Pelagicjuvenile of Pdysiculusjmponictts, 20.4mm SL, FSKU-P 19857, A, Lateral view; B, ventral view of head and abdomen. Seale bars: 5 mm.

anus. Head rounded, dorsum elevated. Mouth moderately large, not protruded; maxilla extending to just behind level of center of eye or slightly beyond. Row of small conical teeth on premaxilla; two or three rows of small conical teeth on den- tary; no teeth on vomer and palatines, Snout rounded with two nostrils, each shorter than eye diameter. Eye large, dorsally situated on head with dorsal margin close to dorsal profile. Interorbital space elevated, width nearly equal to eye diame- ter. Short barbel present on chin. Gill opening wide, opercular region thin. Lumi- nous organ elliptical, situated on midline of belly, nearly at same level as pelvic fin insertion. Narrow line of luminescent tissue beneath skin connecting luminous organ to luminescent area surrounding anus (Fig. 3B). First dorsal fin origin di- rectly above anus; first ray shorter than second ray. Second dorsal fin origin just behind first dorsal fin; last ray not connected to caudal fin by membrane. Anal fin origin a short distance behind anus, approximately below midlength point of first dorsa] fin base. Pectoral fin large, inserted anterior to level of first dorsal fin ori- gin. Pelvic fin with flat base, outer four rays filamentous (ray tips broken). Caudal fin pointed, not confiuent with dorsal or anal fin. Five shallow longitudinal lines on lateral side of body. Scales undeveloped at 20,4mm SL, but appearing on ante- rior trunk at 27,4mm SL, and covering body and head except snout and interor- bital regions at 39.emm SL. Pigmentation in atcohol, Body color mainly light brown, without pigment on any fins. Large melanophores in suborbital and opercular regions, on base of pee- toral fins, and around luminous organ and anus, but luminous organ and lumines- cent area surrounding anus unpigmented (Fig. 3B). Surface of trunk, brain case,

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Pelagic juveniles of t“厂o Pacific morids 23

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and peritoneum covered with sma]1 melanophores. Remarks. Okiyama (1988) described a juvenile of 24,lmm SL referred to Physicutus nzaximowicei Herzenstein, 1896 (a junior synonym of P. jmponicus; see below). It had a luminous organ, six pelvic fin rays, and other meristic characters that supported this identification, but neither the description nor the illustration showed a barbel on the chin. Species of Pltysiculus are characterized by having a small chin barbel (Paulin 1983, 1989b) that appears in the early pelagic juvenile stage (Ambrose 1996; Comyns and Bond 2005), In the present study, even the small- est spectmen (20,4mm SL) had a barbel, as did specimens with parts of the body and head damaged during sampling. Therefbre, Okiyama]s (1988> specimen is un- likely to represent a species of Pdysiculus, In addition, the pigmentation of Okiyama's (1988) juvenile was different from that in our juveniles, with the former specimen having pigmentation on the luminous organ (vs, unpigmented in the present juveniles) and no melanophores on the dorsal half of the trunk (vs. present there in the present juveniles). The lack of a chin barbel and the long anal fin base suggest that Okiyama's specimen is most likely Gadella jordani (BOhlke and Mead, 195I), but the meristic characters do not agree with those of adult G. jordani (see Nakabe 2002). More taxonomic research on the early life stages of Moridae is needed. Although adults of Physicuius J'aponicus are abundant, especially in northern Japan, no specimens in the early life stages, nor even eggs or adults with mature evaries, were collected at depths ranging from 50 to 33em in Iwate Prefecture dur- ing surveys in 1976-1980 (Kitagawa and Nagahora 1983). Kitagawa et at. (1985) re- ported on some moi[phological features and the distribution of eggs and newly hatched larvae of P. jcrponicus based on laboratory-reared and field-caught speci- mens; the latter were collected in March from the inner region of the Kuroshio Current around the Izu Islands. However, Ikeda and Mito (1988) pointed out that the identification of the field-caught eggs and larvae made by Kitagawa et al, (l985) is doubtfUl, owing to the lack of an exact count of myomeres and other characters that distinguish the species. Judging from Kitagawa and Nagahora's (1983) results and the occurrence of pelagic juveniles recorded in this study, P. Jqpontcus may spawn in waters deeper than the continental sheE possibly in the fa11. Physiculus maximotviczi was originally described on the basis of a single speci- men from Hakodate, Hokkaido, northern Japan (Herzenstein 1896), Cohen (1979) and Paulin (1989b) reexamined the holotype and additional specimens and re- garded it as a junior synonym of P. japonicus. In eontrast, Nakabo (2002) recog- nized P. nzaximowiczi as a valid species of PhJ,siculus, indicating the ibllowing characters as diagnostic fbr P. maximowicei: body dark brown (vs. pale brown in P. jcrponicus), eye diameter 2/3 snout length (vs, more than 2/3 snout length in P. jqponicus), and inhabiting depths shallower than 100 m (vs. 150 to 650m in P. jopon- icus). Hewever, the presumed difference in eye diameter can be explained by the frequent protrusien of the eye from the orbital region in fishes brought to the sur- face from great depths. It-is generally well known as well that body color is vari- able according to depth in pelagic fishes; it is highly likely, therefore, that the dif ference in body color noted by Nakabo (2002) is attributable to intraspecific varia- tion and merely reflects depth of habitat. We thus follow Cohen (1979) and Paulin (1989b) and identify the present juveniles as P. J'oponicus.

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25 Pelagic juveniles of two Pacjfic morids

Acknowledgments

We are gratefu1 to Dr Kazuhiro Nakaya (Hokkaido University) for providing valuable information, and Dr Tomio Iwamoto (California Academy of Sciences) and Mr Mark McGrouther (Australian Museum) for their critical reading of the manuscript, Lastly, we thank the captains, otficers, and crew of the R/V Ktz- mamoto-maru for their assistance with sampling.

References

Ambrose, D. A, 1996, Moridae. Pp. 500-507. In: Moser, H, G. (Ed,) The Early Stages ofFtshes in the Calijbrnia C"rrent Region. CalCOFI Atlas, No. 33. Allen Press, Lawrence, Kansas. Cohen, D. M. 1979. Notes on the morid fish genera Lotetla and Physicul"s in Japanese waters. Japanese Journal of lchthyology 26: 225-230. Cohen, D. M. 1984. Gadiformes: overview. Pp. 259-264. tn: Moser, H. G., Richards, W. J., Cohen, D. M., Fahay, M. P., Kendall, A. VV. Jr. and Richardson, S, L. (Eds) Ontogeny and and Her- Systematics ofFishes, Speeial Publication 1. American Society of Ichthyologjsts petotogists, Lawrence, Kansas, Cehen, D. M. 1986. Moridae. Pp. 713-723. ini Whitehead, P. J. P.. Bauchet, M-L,, Hureau, J-C., Nielsen, J. and Tortonese, E. (Eds) Fishes of the Nbrth-eastern Atlantic and the Mediter- ranean, Vbl 2. UNESCO, Paris. J. Early Comyns, B. H. and Bond, P. J. 2005. Moridae. Pp. 617-627. Iit: Richards, W. (Ed.) Atlantic, Stages qf'Attantic Fishes. An ldentij7cation Guide for the IVigstern Central IVbrth Vbl. 1. Tavlor and Francis, New York, San Eschmeyer, W. N. (Ed,) 1998. Catalag of Fishes, Vbts 1 3. Califbrnia Academy of Sciences, Francisco, 2905 pp. Pp, 265-283. Fahav. M. P. and Markle, D. F, 1984. Gadifbrmes: development and relationships. In: Moser, H. G., Richards, W. J., Cohen, D. M., Fahay, M. P., Kendall, A. W, Jr. and Richardson, S. L, (Eds) Ontogeay and Elystematics of Fishes. Special Publication 1. Ameri- can Soctety of Ichthyologists and Herpetologists, Lawrence, Kansas. Fukui, A., Tsuchiya, T. and Uotani, I. 2003. Pelagic juvenile of Hlzlarlv,reus johnsonii (Gadi- formes: Moridae) firom Suruga Bay, Japan. Iehthyological Research 50i 18G-189. Herzenstein, S. M. 1896. Uber einige neue und seltene Fische des Zoologischen Museums der Kaiserlichen Akademie der Wissenschaften. Annuajre del'Academie St. Petersbourg 1: 1-14. M. An Atlas Ikeda, T. and Mito, S. 1988. Pelagic fish eggs. Pp. 999-1083. in: Okiyama, (Ed.) of Press, Tokyo. the Early Stage Fishes in ,Japan, Tokai University [In Japanese] Iwamoto, T. 1975. The abyssal fish Antimora rostrata (Gttnther), Comparative Biochemtstry and Physiology (B) 52i 7-11. Kawai, H. 1972. Hydrography of Kuroshio and Oyashio, Pp. 129-32e. Jn: Iwashita, M, Komaki, Vlol,II. Y,, Hoshino, T., Horibe, S. and Masuzawa, J, (Eds) Physicat Oceanogr(rplzy, Press, Ktityokagak" Kiso Kbza [Fundamental Lecture Qf'Oceanogrqphyl. Tokai University Tokyo. [In Japanese] Kitagawa, D., Kuroda, K. and TsLuruta, Y. 1985. Description and distribution of eggs and lar- of the vae of the brown hakeling Pltysiculus maximowic2i in Japanese waters. Bulletin Japanese Society of Scientific Fisheries 51i 1627-1630. Kitagawa, D. and Nagahora, S. 1983. Estimation of the spawning season ot' the morid fish

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