FISHERIES OCEANOGRAPHY

Fish. Oceanogr. 13:5, 295309, 2004

Differing body size between the autumn and the winterspring cohorts of neon ying squid (Ommastrephes bartramii) related to the oceanographic regime in the North Pacic: a hypothesis

TARO ICHII,1,* KEDARNATH MAHAPATRA,2 MITSUO SAKAI,1 DENZO INAGAKE1 AND YOSHIHIRO OKADA2,3
National Research Institute of Far Seas Fisheries, 5-7-1 Orido, Shimizu, Shizuoka 424-8633, Japan 2 Tokai University Frontier Ocean Research Center (T-FORCE), 3-20-1, Orido, Shimizu, Shizuoka 424-8610, Japan 3 School of Marine Science and Technology, Tokai University, 3-20-1, Orido, Shimizu, Shizuoka 424-8610, Japan
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northward in spring and summer, the autumn cohort has the advantage of being in the SST front and productive area north of the chlorophyll front, whereas the winterspring cohort remains to the south in a less productive area. Thus, the autumn cohort can utilize a food-rich habitat from winter through summer, which, we hypothesize, causes its members to grow larger than those in the winterspring cohort in summer. Key words: neon ying squid, North Pacic, Ommastrephes bartramii, satellite remote sensing, Subtropical Frontal Zone, Transition Zone Chlorophyll Front. INTRODUCTION The neon ying squid, Ommastrephes bartramii, is an oceanic squid occurring worldwide in subtropical and temperate waters (Roper et al., 1984). In the North Pacic, this species plays an important role in the pelagic ecosystem and is an international sheries resource with high commercial value (Seki, 1993). The neon ying squid migrates between its spawning grounds in subtropical waters and feeding grounds in Subarctic waters (Murata and Hayase, 1993). Recently, Polovina et al. (2000, 2001) reported that the Transition Zone Chlorophyll Front (TZCF) migrates seasonally between about 30N in winter and 40N in summer in the north Pacic. The TZCF marks the transition from waters with a low near-surface chlorophyll a (chl a) concentration (0.1 mg m)3) in the south to waters with higher chl a concentrations (0.2 mg m)3) in the north and can be readily monitored by ocean color remote sensing. Albacore tuna (Thunnus alalunga) and loggerhead sea turtles (Caretta caretta) have been shown to exploit the TZCF as a migration pathway and foraging habitat (Polovina et al., 2000, 2001), however, to date it is not clear if or how the life cycle of neon ying squid is associated with the TZCF.
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ABSTRACT The neon ying squid (Ommastrephes bartramii), which is the target of an important North Pacic shery, is comprised of an autumn and winterspring cohort. During summer, there is a clear separation of mantle length (ML) between the autumn (ML range: 38 46 cm) and the winterspring cohorts (ML range: 16 28 cm) despite their apparently contiguous hatching periods. We examined oceanic conditions associated with spawning/nursery and northward migration habitats of the two different-sized cohorts. The seasonal meridional movement of the sea surface temperature (SST) range at which spawning is thought to occur (2125C) indicates that the spawning ground occurs farther north during autumn (2834N) than winter spring (2028N). The autumn spawning ground coincides with the Subtropical Frontal Zone (STFZ), characterized by enhanced productivity in winter because of its close proximity to the Transition Zone Chlorophyll Front (TZCF), which move south to the STFZ from the Subarctic Boundary. Hence this area is thought to become a food-rich nursery ground in winter. The winterspring spawning ground, on the other hand, coincides with the Subtropical Domain, which is less productive throughout the year. Furthermore, as the TZCF and SST front migrate
*Correspondence. e-mail: ichii@affrc.go.jp Received 27 March 2003 Revised version accepted 29 September 2003 2004 Blackwell Publishing Ltd.

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Figure 1. Neon ying squid mantle length variability by month for four putative cohorts (data from Murata, 1990). LL: extra large size group; L: large size group; S: small size group; SS: extra small size group.

Neon ying squid in the North Pacic comprise four size classes, i.e. extra large (LL), large (L), small (S) and extra small (SS) groups, based on monthly mantle length distribution and maturity stages (Fig. 1), as reported by Murata (1990); Yatsu et al. (1997) and Murakami et al. (1981). Unlike the other groups, the LL group consists only of females. During spring and summer, there is a signicant difference in body size between the LL group and the others. For this reason, the LL group was initially believed to live for 2 yr, maturing and spawning in the second year, in contrast to the 1-yr life span of the L and S groups. The SS group was even thought to correspond to the rst year of the LL group (Murata, 1990; Sinclair, 1991). However, recent age estimations from statolith microstructure have shown that all groups have a 1-yr life span, the LL group being an autumn cohort that hatches in September to early January, and the other groups (L, S, SS) forming a winterspring cohort that hatches in January to July (Yatsu et al., 1997). This poses the question as to why such a clear separation of modes in mantlelength composition occurs between the autumn and winterspring cohort despite their contiguous hatching periods (Yatsu et al., 1997, 1998). The size separation could be caused by differences in growth or simply because of separated hatching periods of the two cohorts. Recently Chen and Chiu (2003) demonstrated that the autumn cohort grows faster than the winter cohort, based on intensive examination of statoliths increments. However, there is no report to support the latter possibility. Hence, the size separation can be attributed to the growth difference between two cohorts.

Recently, the spawning locations of the two cohorts were found to differ geographically (Mori et al., 1999). Paralarvae occur at a sea surface temperature (SST) range of about 2125C (Bower, 1994; Mori et al., 1999), suggesting that this is the temperature range at which this species spawns. Based on the seasonal movement of this SST range, Mori et al. (1999) suggested that the autumn cohort spawns farther north (2834N) than the winterspring cohort (2028N). A seasonal shift in the presumed spawning grounds can be seen in Fig. 2 based on the data sets used in Mori et al. (1999). In the present study, we examined the oceanographic regime in this region to investigate possible causes for the size difference between the two cohorts. MATERIALS AND METHODS Data were collected from the spawning grounds, along the northward migration route and at the northern feeding grounds, to examine the inuence of oceanographic variables on these habitats of the neon ying squid. Data were collected from October 1997 through August 1998, when relevant research and shery operations were undertaken. Paralarvae survey The distribution and abundance of paralarvae in the spawning ground were examined from 15 October to 7 November 1997 along ve latitudinal transects (156, 158, 160, 162, and 164W between c. 28 and 37N) at 90-nautical mile intervals. As paralarvae of this species are distributed near the sea surface throughout

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Figure 2. Distribution of paralarval neon ying squid in relation to SST during October, November, February and May. Densities of paralarvae were standardized as numbers of individuals per 20-min surface tow using a ring net (diameter 2 m). The SST range of preferred spawning (2125C; Bower, 1994; Mori et al., 1999) is depicted by bold contours. Paralarval data were pooled over a period of 9 yr (19932001) from the data sets used by Mori et al. (1999). The SST data source is the Reynolds climatological SST database (ftp://ncardata.ucar.edu/datasets/ds277.0/data/oi/mnly/data/).

the day (Young and Hirota, 1990; Saito and Kubodera, 1993; Saito, 1994; Bower, 1996), horizontal surface towing was conducted using a ring net (diameter 2 m,

mesh size 526 lm) at a speed of ca. 2 knots for 10 min. Two tows were carried out at each station. Collected paralarvae of the family Ommastrephidae were

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Figure 3. Distribution of paralarval neon ying squid in relation to (a) SST (C) and (b) surface salinity, during 15 October to 7 November 1997. The boundaries of the Subtropical Frontal Zone (34.6 and 35.2) are shown by the bold salinity contours. For paralarval densities see legend to Fig. 2.

identied to species. A CTD cast was made at each towing station. Drift net survey data Catch data from drift net surveys by the National Research Institute of Far Seas Fisheries (NRIFSF) and Hokkaido University (HU) (Anonymous, 1999a,b) were used to investigate the northward migration of neon ying squid. The NRIFSF survey was conducted along 17000E and 17530E between 35 and 41N during mid- to late-May, 1998, and the HU survey was conducted along 17530E between 40 and 4730N during early August 1998. At each drift net site, 49 or 50 net panels were deployed in the evening and retrieved at the following sunrise. Each panel was 50 m long and 7 m deep. Research drift nets used in the NRIFSF survey comprised 20 commercial mesh nets (stretched mesh size: 115 mm) and 30 non-sizeselective nets [mesh size range (MSR) 48157 mm]. Drift nets used in the HU survey comprised 12 commercial mesh nets (MSR 112118 mm) and 37 non-size-selective nets (MSR 19157 mm). The dorsal mantle length of each neon ying squid was measured to the nearest 1 cm. For those caught in the NRIFSF survey, sex and maturity stages were also determined.

Fishery data Japanese commercial jigging vessels used in the neon ying squid shery ranged in size between 96 and 494 tonnes, and were equipped with about 20100 automatic jigging machines with double reels. To examine the northward migration of neon ying squid, monthly catches derived from commercial shery jigging data were calculated for each 1 1 grid between 30 and 50N, and between 160E and 150W from May through August 1998. A total of 140 shing vessels operated during this season. Satellite image analyses Satellite-derived data were used to investigate SST and chl a concentrations at the spawning grounds during September 1997 to August 1998 and in the northern habitats during May to August 1998. Monthly average SST data were compiled from the Advanced Very High Resolution Radiometer (AVHRR) Pathnder global data sets produced by the National Oceanic and Atmospheric Administration and NASA (Vazquez et al., 1998). Sea-viewing Wide Field-of-view Sensor (SeaWiFS) version 4 space-time binned (level 3) monthly average surface chl a concentration data were

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derived using the global OC4v4 algorithm (OReilly et al., 1998). These products met the 35% accuracy goal xed for SeaWiFS-derived chl a concentrations over a limited, but diverse, set of open ocean validation sites (Hooker and McClain, 2000). To depict the distribution of SST during mid- to late-May and early August 1998, when the drift net surveys were conducted, we estimated the average SST that occurred during May 18 to June 2, 1998 and July 29 to August 13, 1998 using the 8-day average AVHRR Pathnder global SST data sets available during these periods. Likewise, average chl a concentration data during mid- to late-May and early August were derived using the space-time binned, 8-day average SeaWiFS chl a data available for May 9 to June 1, 1998 and July 20 to August 12, 1998, respectively. The SST and chl a data sets were used with spatial resolutions of 0.263 of both latitude and longitude for generating contour lines. Contour lines representing thermal and chl a densities across the study area were computed using a contour plot subroutine of the Generic Mapping Tools (GMT) software package. RESULTS Distribution and abundance of paralarval neon ying squid in autumn Paralarvae of neon ying squid collected in autumn were distributed at SST of 2225C (Fig. 3a). Within this SST range, paralarvae tended to be abundant ( 10 squid/20-min tow) near the surface salinity front, i.e. the Subtropical Frontal Zone (STFZ) dened as the zone occurring between 34.6 and 35.2 isohalines according to Roden (1991), between ca. 29 and 34N (Fig. 3b). The northsouth cross sections along 160, 162 and 164W of salinity and temperature indicated that cooler and less saline water from the north sank below the warmer and higher saline water to the south in the STFZ (Fig. 4). Furthermore, surface salinity fronts extended to ca. 50-m depth. The proximity of paralarvae abundance to the front location suggests that the autumn spawning ground was associated with the STFZ. Oceanographic conditions of spawning/nursery grounds for the autumn and winterspring cohorts of neon ying squid Monthly surface chl a and SST measured by satellite from September 1997 through August 1998 show seasonal changes in oceanographic conditions at the spawning ground (Fig. 5a,b). The autumn cohort spawns in September to early January, and the winter

spring cohort spawns in January to August (Yatsu et al., 1997). Assuming that spawning occurs at 21 25C, the spawning area will shift southward during September to February ahead of the southwardmoving TZCF and then shift northward during March to August behind the northward-moving TZCF. Thus, at no time of the year does the spawning ground occur in the high chl a area. During winter (January to March), the TZCF was located in the southernmost position of the year (Fig. 5a,b), enhancing surface chl a concentrations in the STFZ (2934N). Hence, juveniles of the autumn cohort, which are believed to be distributed in the STFZ, will experience enhanced chl a. However, juveniles of the winterspring cohort never experience enhanced chl a levels because they are believed to be distributed at 2028N (i.e. the Subtropical Domain) where surface chl a was low (< 0.10.2 mg m)3) throughout the year. Thus, the surface chl a levels in the nursery grounds differ between the autumn and winterspring cohorts. Northward migration habitats of neon ying squid in spring and summer Monthly distributions of neon ying squid catches from the jigging shery are shown in relation to various oceanographic fronts from May to August (Fig. 6). Jigging vessels targeted the autumn cohort during these months, because larger individuals (the autumn cohort) have higher commercial value than smaller ones (the winterspring cohort). Over time, the shing areas shifted northward, reaching the Subarctic Boundary in June and the Subarctic Front in August. Compared with the monthly location of the TZCF, substantial catches were taken north of the chlorophyll front in areas where surface chl a concentrations were higher than 0.20.3 mg m)3. The seasonal northward shift of shing areas also coincided with the northward shift of SST ranging from 12 to 18C (the SST front). Mantle length (ML) compositions of neon ying squid caught by non-size-selective drift net surveys shows that larger individuals were distributed farther north than smaller individuals (Fig. 7). Mature individuals caught in May were considered to be southward migrating members of the winterspring cohort, so they were excluded from the following analysis. Individuals were divided into the northward migrating autumn (i.e. ML 30 cm for May and 34 cm for August) and winterspring cohorts (ML < 30 cm for May and <34 cm for August) according to Murata and Hayase (1993). The low number of the winterspring cohort

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Figure 4. Density of paralarval neon ying squid in relation to northsouth sections of salinity and temperature (C) along three latitudinal transects (a)164W, (b)162W and (c)160W, during 15 October to 7 November 1997. The Subtropical Frontal Zone corresponds to the area between the bold salinity contours 34.6 and 35.2 at the surface. Horizontal bars at the top indicate the Subtropical Frontal Zone. ., m: station locations. For paralarval densities see Fig. 2.

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Figure 5. (a) Monthly satellite images of SeaWiFS chl a (mg m)3) (left panels) and AVHRR-derived SST (C) (right panels), during September 1997 to February 1998. Bold line in left panels indicates the Transition Zone Chlorophyll Front. SST of spawning preference (2125C) is depicted by bold contours. (b) Monthly satellite images of SeaWiFS chl a (mg m)3) (left panels) and Advanced Very High Resolution Radiometer (AVHRR)-derived SST (C) (right panels), during March to August 1998. Bold line in left panels indicates the Transition Zone Chlorophyll Front. SST of spawning preference (2125C) is depicted by bold contours.
(a)

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Figure 5. Continued.
(b)

caught in May suggests that a substantial part of the winterspring cohort was distributed south of the survey area at that time. Catch compositions of the drift net surveys indicated that the autumn and winter

spring cohorts were separated by the TZCF during both May and August (Fig. 8); the winterspring cohort occurred in low surface chl a ( 0.2 mg m)3) areas and the autumn cohort occurred in high chl a areas

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Figure 6. Monthly distribution of neon ying squid catches (tonnes) from the Japanese jigging shery for each 1latitude 1 longitude in relation to SeaWiFS chl a (mg m)3) (left panels), satellite-derived SST (C) (middle panels) and water temperature at 120 m depth (C) (right panels), during May to August 1998. The bold line in the left panels indicates the Transition Zone Chlorophyll Front. The 4 and 8C isotherms at 120 m depth in the right panels indicate the Subarctic Front and Subarctic Boundary, respectively (Uehara, 1992). Temperature data at 120 m are from the Joint Environmental Data Analysis Center (http://jedac.ucsd.edu/DATA_IMAGES/ index.html).

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Figure 7. Mantle length composition of neon ying squid caught with non-selective drift nets at each station along (a) 17000E and 17530E in mid- to late-May and (b) 17530E in early August, 1998. Combined frequencies from all stations during the respective periods are shown at the bottom. MI: immature males; MM: mature males; FNC; non-copulated females; FC: copulated females.

(> 0.2 mg m)3) during their northward migration. The two cohorts were also separated by a zone with an SST range of 1618C. While the autumn cohort occurred in association with the SST front, the winterspring cohort occurred mainly south of it. DISCUSSION Oceanographic factors related to the formation of neon ying squid spawning grounds in autumn Neon ying squid paralarvae were abundant in the STFZ within the SST range of spawning preference

between 21 and 25C in autumn (Figs 3 and 4). In a survey undertaken by the rst author in autumn 2001 in the same area, paralarvae were abundant in the STFZ within the same SST range. Ishino (1975) suggested that oceanic fronts play an important role in the formation of sh concentrations in three ways: (i) by forming a barrier to sh movement, (ii) through the accumulation of surface drifting food, and (iii) by maintaining high productivity. Strong shear between the STFZ and Subtropical Domain (Seki et al., 2002) may have acted as a barrier to movement, facilitating the accumulation of spawning squid.

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Figure 8. Distribution of neon ying squid CPUE (number of individuals per 30 non-size-selective drift nets) in relation to SeaWiFS chl a (mg m)3) (left panels) and AVHRR-derived SST (C) (right panels) for mid- to late-May and early August 1998. Autumn and winterspring cohorts shown by black and white circles, respectively. Denitions of the cohorts are indicated in text. * in upper panels indicates the jigging station with no squid catch. Bold line in left panels indicates the Transition Zone Chlorophyll Front.

Spawned egg masses of ommastrephid squids are presumed to be suspended around the depth of the pycnocline (thermo- and/or haloclines) and require water temperatures 14C for successful embryonic development (ODor et al., 1982; ODor and Balch,1985; Sakurai et al., 1996; Sakurai, 2002). Oceanographic conditions in the STFZ met the temperature requirement for embryonic survival, as temperatures in the pycnocline were well above 14C (Fig. 4). The STFZ occurs in a region between the westerly (to the south) and the easterly (to the north) trade winds, so it is a zone of Ekman transport convergence in the central north Pacic (Roden, 1975). Drifters released in the STFZ were reported to remain near the region (Roden, 1991), suggesting paralarvae of the

autumn cohort that hatch here will remain in the zone, which becomes productive but colder during winter. Contrasts in productivity of the spawning/nursery grounds for the autumn and winterspring cohorts of neon ying squid The STFZ spawning ground of the autumn cohort was characterized by enhanced surface chl a concentrations in winter because of the proximity of the TZCF, which moved southward to the STFZ in winter from its summer location at the Subarctic Boundary (Fig. 5a,b). Recent works have identied two prominent winter features of the STFZ, the Subtropical Front (STF) and South Subtropical Front (SSTF) (Polovina et al., 2000; Leonard et al., 2001; Seki et al.,

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Figure 9. CPUE (number of individuals per 30 non-size-selective drift nets) for shes and squids collected along 17530E in (a) mid- to late-May and (b) early August 1998. Black and white parts of the circles represent the autumn and the winterspring cohorts, respectively. (The small mesh sizes of 1938 mm of drift net were not used during May, so small sh species, such as Japanese anchovy and Pacic saury, were not collected.)

(a)

(b)

2002). The STF occurs near the 17C SST isotherm and coincides with the TZCF, and the SSTF occurs near the 21C SST and coincides with an increase in subsurface chl a concentration. Thus, integrated chl a concentration in the water column is enhanced in the STFZ during winter (Leonard et al., 2001; Seki et al., 2002). The STFZ becomes a good shing ground for swordsh (Xiphias gladius) during winter and spring (Seki et al., 2002). Research surveys suggest that its main prey, i.e. neon ying squid, is most abundant in the STFZ, although the spawning ground of the squid at this time of year occurs farther south (Young and Hirota, 1999; Seki et al., 2002). Furthermore, stomach content data for neon ying squid suggest that the number of sh eaten (estimated from otoliths and eye lenses) increases near the STFZ (Young and Hirota, 1999). Therefore, the increased biological productivity associated with the STFZ in winter may provide the framework for a food-rich nursery ground for juveniles of the autumn cohort. On the contrary, the spawning ground of the winter spring cohort, i.e. the Subtropical Domain, is less productive throughout the year (Fig. 5a,b). The stable subtropical surface layer that generally occurs in the upper 100 m (Roden, 1991) is the principal force inhibiting the vertical ux of nutrients into the euphotic

zone (Mann and Lazier, 1991). In fact, uxes of nutrients into the euphotic zone in the Subtropical Domain are probably the lowest of any oceanic environment (Cullen, 1982). Thus, feeding conditions may be poor in the nursery ground of the winterspring cohort. Forsythe (1993) suggested the importance of water temperature for eld growth of young squid; a small (12C) increase in water temperature experienced by exponentially growing squid during the rst 24 months after hatching can produce signicantly faster growth. In the case of neon ying squid in the North Pacic, the autumn cohort experiences colder temperatures (1620C) than the winterspring cohort (2125C) during the nursery period because the former occurs farther north than the latter. Considering that the autumn cohort grows faster than the winterspring cohort during the juvenile and subadult stages, food conditions may have relatively more effect on the growth of neon ying squid than water temperature does, as in the case of the California market squid Loligo opalescens (Jackson and Domeier, 2003). Contrasts in northward migration habitats of neon ying squid in spring and summer As the TZCF shifted northward in spring and summer, the autumn cohort had the advantage of being in the SST front and high surface chl a concentrations

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(> 0.2 mg m)3) north of the TZCF (Figs 6 and 8), whereas the winterspring cohort remained in a low surface chl a area (0.2 mg m)3) to the south (Fig. 8). Watanabe et al. (2004) showed that stomach fullness of the autumn cohort tended to be greater than in the winterspring cohort during their northward migration. Furthermore, according to the drift net surveys of this study, many epipelagic nekton, such as Pacic pomfret (Brama japonica), Pacic saury (Cololabis saira), Japanese anchovy (Engraulis japonicus), blue shark (Prionace glauca) and boreal clubhook squid (Onychoteuthis borealijaponicus) also forage north of the TZCF, as in the case of the autumn cohort (Fig. 9). Only skipjack tuna (Katsuwonus pelamis) does so south of the front. Hence, from a trophic standpoint, the higher chl a area north of the TZCF may set the framework for an enhanced feeding regime for many epipelagic nekton, compared with the low chl a area to the south. The diet of the autumn cohort includes shes (mainly myctophids, followed by anchovy and Pacic saury) and squids (such as the boreal clubhook squid, Onychoteuthis borealijaponica, and the minimal armhook squid, Berryteuthis anonychus) in spring and summer (Naito et al., 1977; Sinclair, 1991; Watanabe et al., 2004). Many of these prey species, such as anchovy, Pacic saury, boreal clubhook squid and some myctophid shes are migratory (Ogawa, 1961; Kubodera, 1986; Shimazaki, 1986). Hence, it appears the autumn cohort follows these migratory shes and squids in their northward migration. The winter spring cohort, on the other hand, feeds on euphausiids, amphipods and small sh species, such as Maurolicus imperatorius (Watanabe et al., 2004), which are considered to be endemic to the Transition Zone. Thus, there appears to be a marked contrast in the food webs between the high chl a area north of the TZCF and low chl a area to the south. Overall food availability for neon ying squid The habitat of juveniles and subadults of the autumn cohort appears to be more food rich than those of the winterspring cohort. This could be one reason why there is a clear separation, during summer (Fig. 7b), in ML between the autumn cohort (ML range: 38 46 cm; mode: 43 cm) and the winterspring cohort (ML range: 1628 cm; modes: 18, 22, and 26 cm) in spite of no apparent separation of hatching periods between them. Females of the winterspring cohort, however, compensate for their slow growth by penetrating the Subarctic Boundary and moving northward into the Transitional Domain, feeding there during summer and fall. Consequently, by the time of

spawning their nal body size equals that of the autumn cohort (Fig. 1; Murata and Hayase, 1993). Whereabouts of males of the autumn cohort are unknown but speculated to be in the southern Transition Zone during spring and summer, when females occur in the northern Transition Zone and Transitional Domain (Yatsu et al., 1997). In the case of the winterspring cohort, immature males and females occur in equal numbers until they migrate to the Subarctic Boundary (Kubodera et al., 1983; Murata and Nakamura, 1998). At this point, only females penetrate into the northern area of the Transitional Domain, most males remaining around the Subarctic Boundary. This suggests that, whether or not they use a food-rich area, e.g. the Transitional Domain, during feeding, migration may lead to an extraordinary difference in size between the sexes (Fig. 1). ACKNOWLEDGEMENTS We thank J. Mori for sampling and identication of paralarval neon ying squid. We are also grateful to J. R. Bower and H. Watanabe for their critical readings of the manuscript. Our sincere thanks are due to the captains, ofcers and crew of the R.V. Wakatori Maru for their assistance during the cruise. The authors would like to thank the SeaWiFS Project (Code 970.2) and the Goddard Earth Sciences Data and Information Services Center/Distributed Active Archive Center (Code 902) at the Goddard Space Flight Center, Greenbelt, MD 20771, for the production and distribution of SeaWiFS data, respectively. These activities are sponsored by NASAs Earth Science Enterprise. REFERENCES
Anonymous (1999a) Data Record of Oceanographic Observations and Exploratory Fishing. Hakodate, Japan: Faculty of Fisheries, Hokkaido University, No. 42. Anonymous (1999b) Data Record of Oceanographic Observations and Exploratory Fishing. Hakodate, Japan: Faculty of Fisheries, Hokkaido University, No. 42 (Supplement). Bower, J.R. (1994) Distribution of paralarvae of the squid Ommastrephes bartramii near the Hawaiian archipelago. Masters Thesis, Department of Oceanography, University of Hawaii, 120 pp. Bower, J.R. (1996) Estimated paralarval drift and inferred hatching sties for Ommastrephes bartramii (Cephalopoda:Ommastrephidae) near the Hawaiian Archipelago. Fish. Bull. 94:398411. Chen, C. and Chiu, T. (2003) Variations of life history parameters in two geographical groups of the neon ying squid, Ommastrephes bartramii, from the North Pacic. Fish. Res. 63:349366.

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