FISHERIES OCEANOGRAPHY
24:2, 190–203, 2015
Seasonal potential fishing ground prediction of neon flying
squid (Ommastrephes bartramii) in the western and central
North Pacific
IRENE D. ALABIA,1* SEI-ICHI SAITOH,1
ROBINSON MUGO,2 HIROMICHI IGARASHI,3
YOICHI ISHIKAWA,3 NORIHISA USUI,4
MASAFUMI KAMACHI,4 TOSHIYUKI AWAJI5
AND MASAKI SEITO6
1
Laboratory of Marine Environment and Resource Sensing,
Graduate School of Fisheries Sciences, Hokkaido University,
3-1-1 Minato-cho, Hakodate, Hokkaido 041-8611, Japan
2
Regional Centre for Mapping of Resources for Development -
SERVIR-Africa Kasarani Road, P.O. Box 632-00618,
Ruaraka, Nairobi Kenya
3
Data Research Center for Marine-Earth Sciences, Japan Agency
for Marine Earth-Science and Technology (JAMSTEC), 3173-
25 Showamachi, Kanazawa-Ward, Yokohama City, Kanagawa
236-0001, Japan
4
Oceanographic Research Department, Meteorological Research
Institute, 1-1 Nagamine, Tsukuba, 305-0052, Japan
5
Graduate School of Science Division of Earth and Planetary
Science, Kyoto University, Sakyo-Ward, Kyoto 606-8502,
Japan
6
Aomori Prefectural Industrial Technology Research Center,
4-11-6 Dainitonya-Machi, Aomori-Shi, Aomori 030-0113,
Japan
ABSTRACT
We explored the seasonal potential fishing grounds of
neon flying squid (Ommastrephes bartramii) in the wes-
tern and central North Pacific using maximum entropy
(MaxEnt) models fitted with squid fishery data as
response and environmental factors from remotely
sensed [sea surface temperature (SST), sea surface
height (SSH), eddy kinetic energy (EKE), wind stress
curl (WSC) and numerical model-derived sea surface
salinity (SSS)] covariates. The potential squid fishing
grounds from January–February (winter) and June–July
(summer) 2001–2004 were simulated separately and
covered the near-coast (winter) and offshore (summer)
*Correspondence. e-mail: irenealabia@salmon.fish.hokudai.
ac.jp
Received 24 June 2013
Revised version accepted 14 January 2015
190 doi:10.1111/fog.12102
Fish. Oceanogr.
forage areas off the Kuroshio–Oyashio transition and
subarctic frontal zones. The oceanographic conditions
differed between regions and were regulated by the
inherent seasonal variability and prevailing basin
dynamics. The seasonal and spatial extents of poten-
tial squid fishing grounds were largely explained by
SST (7–17°C in the winter and 11–18°C in the sum-
mer) and SSS (33.8–34.8 in the winter and 33.7–34.3
in the summer). These ocean properties are water mass
tracers and define the boundaries of the North Pacific
hydrographic provinces. Mesoscale variability in the
upper ocean inferred from SSH and EKE were also
influential to squid potential fishing grounds and are
presumably linked to the augmented primary produc-
tivity from nutrient enhancement and entrainment of
passive plankton. WSC, however, has the least model
contribution to squid potential fishing habitat relative
to the other environmental factors examined. Findings
of this work underpin the importance of SST and SSS
as robust predictors of the seasonal squid potential fish-
ing grounds in the western and central North Pacific
and highlight MaxEnt’s potential for operational fish-
ery application.
Key words: maximum entropy model, neon flying
squid, North Pacific, potential fishing ground, satellite
data
INTRODUCTION
Neon flying squid (Ommastrephes bartramii) is a large
pelagic cephalopod distributed in the temperate and
subtropical waters worldwide (Roper et al., 1984). It is
among the economically important fisheries exploited
internationally in the North Pacific. Japan has com-
mercially harvested it since 1974, concurrent with the
period when the abundance of the Japanese common
squid Todarodes pacificus plummeted (Yatsu et al.,
2000). The resource harvesting operations were later
joined by South Korea, Taiwan and China (Yatsu
et al., 1997). The Japanese squid driftnet fishing sea-
son begins in June and lasts to December and 50% of
the average annual catch since 1983 occurred in July
© 2015 John Wiley & Sons Ltd
 Seasonal potential fishing ground prediction of neon flying squid
and August (Ignell, 1991). From 1996 to 2000, the
fishing season began earlier, towards the end of April
and the beginning of May (Liu and Chen, 2002) and
extended to November (Chen and Tian, 2006).
The North Pacific population of O. bartramii is
comprised of the fall and winter–spring spawning
cohorts and makes a clock-wise, annual round-trip
migration between its spawning grounds (subtropical
region) and its northern feeding grounds (subarctic
boundary and transitional domain) (Murata and
Nakamura, 1998). The primary migratory route is
thought to be related to the flow of the northern
warm-water branches of the Kuroshio Current and its
associated frontal zone (Shao et al., 2005). Addition-
ally, the life history differences between the two sea-
sonal spawning cohorts have been attributed to
oceanographic regime shifts in the optimum spawning
and feeding zones (Ichii et al., 2009). The neon flying
squid also undergoes a south–north feeding migration
between the subtropical region and the transitional
domain north of the subarctic boundary as the latter is
highly productive in summer (Watanabe et al., 2004).
The changes in ommastrephid squid abundance
are to a larger extent, influenced by the growing glo-
bal fishing pressure and climatic shifts (Waluda et al.,
2001; Waluda and Rodhouse, 2006). Short-lived spe-
cies such as cephalopods have been continually
increasing as a proportion of marine capture fisheries
and maintain or increase the world capture fisheries
production (Chen et al., 2008). The short-life span
and semelparity of cephalopods also make them
highly vulnerable to drastic changes in environmen-
tal conditions (Rodhouse, 2001). The increasing
influence of externalities in population dynamics of
marine resources necessitates the utilization of robust
tools to accurately deduce the extent and geographi-
cal distribution of fisheries for conservation and man-
agement. The occurrence and abundance of the neon
flying squid in the western North Pacific are affected
by its environment. Previous studies have shown that
sea surface temperature (SST) is the primary environ-
mental variable influencing the distribution of squid
fishing grounds (Cao et al., 2009; Tian et al., 2009a).
However, other oceanographic and atmospheric fac-
tors including sea surface salinity (SSS), mesoscale
variability and wind stress curl (WSC) could directly
or indirectly impact fish habitat. SSS is known to
affect the distribution of the neon flying squid
habitat, and its spawning ground is related to surface
salinity fronts (Ichii et al., 2004). Mesoscale struc-
tures and associated variability also account for neon
flying squid abundance (Watanabe et al., 2008; Chen,
2010), and WSC is known to influence the formation
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
191
of these structures and frontal systems in the western
North Pacific (Kappes et al., 2010). In this study, the
effects of environmental variables (including less
explored variables such as EKE and WSC) to squid’s
potential fishing grounds in the western and central
North Pacific was examined using a statistical model-
ing approach.
Habitat suitability modeling coupled with geo-
graphic information system (GIS) technology has
been widely used to deduce the distribution of fish spe-
cies in relation to environmental factors (Manderson
et al., 2011; Gong et al., 2012). However, many of the
existing statistical modeling methods require compre-
hensive and accurate presence and absence data that
are rarely available especially for highly mobile marine
species. It has been suggested that the use of presence-
absence data results in superior model performance
and reliable predictions (Hirzel et al., 2001). How-
ever, incorrect assessment of absence data, geographi-
cal sampling biases, and inadequate sampling
procedures also reduce the performance and accuracy
of model predictions (Pearson et al., 2007; Jones et al.,
2012). Thus, alternative presence-only approaches are
of special interest in model construction, aimed at
generating habitat suitability maps for marine species
(MacLeod et al., 2008; Ready et al., 2010).
Maximum entropy (MaxEnt) (Phillips et al., 2006)
is a machine learning method that is widely used in
the terrestrial and marine habitat modeling realm.
The MaxEnt algorithm uses a generative approach to
estimate the effects of environmental layers to species’
presence by approximating an unknown probability
distribution with maximum entropy subject to con-
straints that represent incomplete information. Recent
studies have shown that MaxEnt satisfactorily pre-
dicted the potential habitat of cephalopods and estab-
lished their habitat responses to oceanographic
conditions in the Atlantic Ocean (Lefkaditou et al.,
2008; Hermosilla et al., 2011). In the western North
Pacific, regression-based and mean-field models were
widely utilized to predict squid’s suitable habitats using
fishery-dependent abundance data (Tian et al., 2009b;
Chen et al., 2011). However, presence-only models
such as MaxEnt remain less explored, and implemen-
tation of this approach could be useful for predicting
the neon flying squid potential fishing habitats and
understanding squid–environment interactions. Our
objectives were (i) to assess the effects of environmen-
tal parameters to potential squid fishing grounds; (ii)
to examine seasonal potential fishing grounds of
O. bartramii in the western and central North Pacific;
and (iii) to evaluate the performance of MaxEnt for
squid fishing ground prediction.
192 I. D. Alabia et al.
MATERIALS AND METHODS
Study area
The study area covers the western North Pacific
(140°E–160°W and 30°N–50°N) (Fig. 1), where the
Japanese squid-jigging operation extends off the east-
ern coast of Japan towards the offshore waters. It is an
extensively dynamic and productive ecosystem mainly
influenced by the warm Kuroshio Current and the cold
Oyashio Current (Talley, 1993). The Kuroshio
originates from the warm and saline current of the
Subtropical Gyre (Talley et al., 1995) whereas the Oy-
ashio forms from the cold and less-saline waters of the
Western Subarctic Gyre (Yasuda, 2003). The conflu-
ence of the two predominant western boundary cur-
rents flows between the subtropical and subarctic
frontal zones forming the Kuroshio–Oyashio transition
zone. The zone is characterized by mesoscale energetic
features and formation of various water masses (Roden,
1991).
Squid fishing locations
The squid occurrence records used to fit the habitat
suitability models (HSM) were obtained from the daily
squid fishing locations from December–February and
May–July of 2001, representative of periods with the
highest Japanese jigging activities (Fig. 3a). These
were provided by the Aomori Prefectural Industrial
Technology Research Center (APITRC). The daily
fishing logs (point data) with information on fishing
date (Month, Day and Year) and locations (Longitude
and Latitude) were compiled into monthly databases
prior to model construction. The independent sets of
squid fishing logs from January–February and June–July
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
2002–2004 were used for model validation. To mini-
mize the data inconsistencies in the model validation,
December and May were no longer included owing to
unavailability of data in December 2004 and only a
few data samples in May.
Environmental data
Satellite-derived measurements of SST, sea surface
height (SSH), EKE, WSC and numerical model-
derived sea surface salinity (SSS) from 2001 to 2004
were used as the environmental predictors for the sta-
tistical models. Daily SST data were compiled from
the Level 4 optimally-interpolated (OI) Advanced
Very High Resolution Radiometer (AVHRR) SST
version 2, generated by the Physical Oceanography
Distributed Archive Center (PO.DACC) at the
NASA Jet Propulsion Laboratory (JPL) (ftp://podaac-
ftp.jpl.nasa.gov/allData/ghrsst/data/L4/GLOB/NCDC/
AVHRR_OI/) at 0.25° resolution. The daily NASA
QuikSCAT (Quick Scatterometer)-derived WSC at
0.125° spatial resolution was downloaded from NOAA
Environmental Research Division’s Data Access Pro-
gram (ERDDAP) (http://coastwatch.pfeg.noaa.gov).
The daily delayed-time, updated and merged product
of mapped absolute dynamic topography (MADT) for
SSH and the merged product of the mapped sea level
anomaly (MSLA) geostrophic velocities (u,v) were
used to compute the EKE provided by the Archiving
Validation and Interpretation of Satellite Data (AVI-
SO) at a 0.33 9 0.33° spatial resolution on a Mercator
grid (ftp://aviso.oceanobs.com). The monthly averages
for each of the parameters were computed from the
daily available datasets using bourne again shell (bash)
scripts. EKE measurements were computed from
Figure 1. Map of the study area showing
the hydrographic and topographic fea-
tures of the basin. Dashed white line
divides the study site into two regional
model domains (Region 1:140–150°E and
Region 2:150°E-160°W) based on dis-
tinct clusters of squid fishing points (filled
circles) from 2001 to 2004. The model
predictions of the potential squid fishing
habitat were mapped in areas denoted by
the dashed black lines.
200 I. D. Alabia et al.

Figure 8. The spatial distribution of squid fishing locations (colored dots) and predicted ranges of sea surface temperature (SST)
(solid lines) and sea surface salinity (SSS) (dashed lines) for June and July 2001–2004 overlain on habitat suitability maps pro-
jected using 2001 regional models.

Table 2. Computed area under the curve (AUC) from were sufficiently sampled in base models). MaxEnt
monthly models for January–February and June–July 2002– models, however, were unable to satisfactorily extrapo-
2004 projected using the 2001 base models. AUCs for the late the squid fishing grounds across regions with dif-
2001 base models are presented in bold. ferent environmental conditions from the base
AUC models. Thus, MaxEnt’s utility for operational fishery
applications (i.e., higher prediction accuracy) could
Model 2001 2002 2003 2004 probably be improved by circumventing data and sam-
January 0.942 0.963 0.756 0.742 pling limitations.
February 0.984 0.801 0.846 0.841 Finally, model contributions of environmental vari-
June 0.983 0.804 0.822 0.756 ables used to predict potential squid fishing habitat
July 0.969 0.793 0.589 0.847 will be discussed. In all the regional models, SST and
SSS were the most important oceanographic factors
affecting the potential fishing habitat, followed by
sampling bias in this study, MaxEnt models showed a SSH, EKE and WSC (Fig. 5). The high primary SST
strong predictive ability (i.e. based on the statistical and SSS model contributions are consistent with find-
performance (Tables 1 and 2) and modest spatial cor- ings of earlier studies showing their robust influence
respondence (Figs 7 and 8) in predicting seasonal on O. bartramii distribution and abundance in the
potential squid fishing grounds at least in sites that North Pacific (Yatsu and Watanabe, 1996; Bower and
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
194 I. D. Alabia et al.
Figure 2. Spatial distribution of squid occurrence data overlaid on images of (a) sea surface temperature (SST), (b) sea surface
salinity (SSS), (c) sea surface height (SSH), (d) eddy kinetic energy (EKE) and (e) wind stress curl (WSC) from January–Febru-
ary and June–July 2001–2004.
(a)
(b)
(c)
(d)
(e)
from 2001 to 2004 are shown in Fig. 3a–e. The squid
jigging activities showed an increase in operation dur-
ing the summer relative to the winter season (Fig. 3a)
possibly owing to restriction imposed by poor weather
conditions during winter and consideration of the
existing knowledge of seasonal timing of squid migra-
tion. The mean SST showed a lower temperature of
the fishing sites in winter relative to the summer sea-
son (Fig. 3a) and reflects seasonal temperature
changes. The mean SSS distribution (Fig. 3c) was
highest from January to February and May to June
when fishing points were primarily concentrated along
the Kuroshio–Oyashio transition zone (35–40°N).
The mean SSS correspondingly dropped in July as the
jigging activities shifted north towards the Subarctic
Frontal Zone (SAFZ) around 41–45°N. The mean
SSH (Fig. 3d) distribution of the squid fishing posi-
tions showed downward trends for winter and summer.
During winter, the decrease in mean SSH reflects a
seasonal cooling signal. However, an observed down-
ward trend in summer corresponds to the northward
shift of jigging fleets to SAFZ. The mean EKE (Fig. 3e)
during winter showed an increasing trend correspond-
ing to the southward shift of the jigging fleets closer to
the Kuroshio Current. During summer, however, the
mean EKE declined from May to July as the squid jig-
ging fleets moved farther east, away from the influence
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
of strong western boundary currents. From May to
June, the mean EKE showed modest values
(≥100 cm2 s2) when squid fishing fleets operate along
the eastern part of the Kuroshio–Oyashio transition
zone. The mean WSC during the winter showed a
stronger negative curl relative to the summer season
when the winds are relatively calmer indicated by
near-zero mean WSC (Fig. 3f).
Squid distribution in relation to environmental variables
The distributions of squid fishing locations in the 4-yr
period in relation to environmental parameters in the
western North Pacific for January–February and June–
July are shown in Fig. 4a–e and f–j, respectively. In Janu-
ary–February (Region 1), squid fishing activities
occurred in regions with a SST between 7–17°C and
were highest at 9°C (Fig. 4a). The SSS ranged from 33.8
to 34.8 with highest fishing activity at 34.4 (Fig. 4b).
The SSH ranged from 15 to 75 cm with the highest fish-
ing activity at 35 cm (Fig. 4c). The EKE ranged
between 50–1000 cm2 s2 with a peak at 250 cm2 s2,
while WSC ranged from 1.5 to 1.5 MPa m1 with a
peak at 0.25 MPa m1. In June–July (Region 2), the
SST ranged from 11 to 18°C with a peak at 13°C
(Fig. 4f); SSS from 33.1 to 34.5 with peaks at 33.7 and
34.3 (Fig. 4g); SSH from 25 to 75 cm with peak at
55 cm (Fig. 4h); EKE between 0–300 cm2 s2 with
 Seasonal potential fishing ground prediction of neon flying squid 195

Figure 3. Monthly mean time-series of (a) number of squid fishing locations, (b) sea surface temperature (SST), (c) sea surface
salinity (SSS), (d) sea surface height (SSH), (e) eddy kinetic energy (EKE) and (f) wind stress curl (WSC) from 2001 to 2004
for the winter (December-February) and summer (May-July). The vertical bars correspond to  1 SD.

(a) (b)

(c) (d)

(e) (f)

peak at 50 cm2 s2 (Fig. 4i) and WSC from –0.4 to
0.25 MPa m1 with peak at 0.1 MPa m1 (Fig. 4j).
MaxEnt model performance and squid’s potential habitat
characteristics
The results of MaxEnt models showed that all the
monthly regional models performed significantly bet-
ter than random with AUC > 0.9 (CI = 99)
(Table 1). Model performance statistics for May and
December, although better relative to the other
months, were not considered owing to relatively low
sample sizes and unavailability of test data for evalua-
tion. Based on the computed model statistics, January–
February and June–July were selected as representative
models for winter and summer periods, respectively.
The contribution of the environmental parameters to
model prediction is shown in Fig. 5. Results showed
that SST, SSS, SSH and EKE were among the envi-
ronmental predictors with the highest contribution to
the model gain. Among the four variables, SST and
SSS mostly accounted for the corresponding increase
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
in model gain. WSC recorded the least contribution in
all regional models developed for O. bartramii.
Model-derived preferred ranges for the most impor-
tant environmental variables (SST, SSS, SSH and
EKE) are shown in Fig. 6. In January, the derived
environmental ranges in squid potential habitat
were between 7–15°C, 33.4–34.4, 25–60 cm and
0–3000 cm2 s2 for SST, SSS, SSH and EKE, respec-
tively. In February, SST, SSS, SSH and EKE
ranged from 7–14°C, 33.6–34.5, 15–50 cm and
0–1000 cm2 s2, respectively. During these periods,
squid fishing fleets were located in regions of colder
SST ranges with a higher surface salinity relative to
the summer months (June–July). Similarly, winter fish-
ing locations (January-February) were characterized by
a lower SSH and higher EKE ranges. In June and July,
the model-derived ranges for SST and SSS were from
10–17°C and 33.5–34.3 and 12–18°C and 33.0–34.0,
respectively. In June, SSH and EKE ranged from 40–
80 cm and 0–1250 cm2 s2 and dropped to 30–60 cm
and 0–100 cm2 s2 in July.
196 I. D. Alabia et al.

(a) (b) (c)

(d) (e)

(f) (g) (h)

(i) (j)

© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
 Seasonal potential fishing ground prediction of neon flying squid 197

Figure 4. Frequency distributions of sea surface temperature (SST) (a, f), sea surface salinity (SSS) (b, g), sea surface height
(SSH) (c, h), eddy kinetic energy (EKE) (d, i) and wind stress curl (WSC) (e, j) extracted using the available squid fishing
records from January–February and June–July 2001–2004, respectively.

extended towards the offshore waters in February with

Prediction and validation of Ommastrephes bartramii
occurrence
Mapped habitat suitability indices (HSI) for selected
base models (2001) and predicted HSI for similar
monthly periods (2002–2004) are shown in Figs 7 and
8, respectively. In representative winter months
(Fig. 7), the predicted probability of squid occurrence
covered the areas adjacent to the coast in January and
Table 1. Summary statistics derived from monthly models
for representative summer (May–July) and winter (Decem-
ber–February) months of 2001. Base models were calibrated
using 70% of occurrence points and area under the curve
(AUC) and SD were calculated from the remaining 30% of
the presence records. Total number of fishing locations for
each month is denoted by N. All models fit significantly bet-
ter than chance (AUC > 0.90) at CI = 99%.
high probability zones (HSI ≥ 0.6) between 36–41°N.
In summer period (Fig. 8), the predicted HSI extended
across the oceanic waters of the western North Pacific.
In June, squid’s potential habitats were distributed
across 165°W–160°E and 38–42°N. The predicted
probability of occurrence slightly shifted north in July
(40–45°N) and the northern boundaries (42–43°N)
exhibited higher habitat probability values relative to
the southern boundaries (40–41°N). The predicted
HSI distribution for January–February and June–July
2002–2004 generally showed modest spatial correspon-
dence with actual squid fishing locations albeit spatial
mismatches were particularly high for January and July
2003 predictions. The extent of the squid potential
fishing habitat was bounded by SST and SSS ranges
derived from monthly selected base models.

MaxEnt models Regional domain N AUC SD DISCUSSION

January-01 1 200 0.942 0.032 We utilized squid fishing locations and oceanographic
February-01 1 171 0.984 0.004 variables to predict the seasonal potential fishing
May-01 2 83 0.974 0.006 grounds of O. bartramii in the western and central
June-01 2 432 0.983 0.003 North Pacific using MaxEnt models. While species
July-01 2 361 0.969 0.007 occurrences from fisheries-dependent datasets suffer
December-01 1 242 0.961 0.009
from sampling bias introduced through behavioral

Figure 5. Jacknife test results of variable

importance derived for the regional mod-
els. The gray and white bars correspond
to the effect on model gain by using and
sequentially removing each variable from
the model, respectively.

© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
198 I. D. Alabia et al.

Figure 6. Response curves for important variables (a) sea surface temperature (SST), (b) sea surface salinity (SSS), (c) sea sur-
face height (SSH) and (d) eddy kinetic energy (EKE) in all regional models. Solid lines correspond to the effect of each variable
on the model; dashed lines represent the 99% confidence interval.

(a) (b) (c) (d)

effects related to fishing operations, economic and
government restrictions, this information is inexpen-
sive, most abundant and widely available for fisheries
scientists and managers (Gallaway et al., 2003; Mugo
et al., 2010; Dell et al., 2011). An implicit assumption
in utilizing these data is that regions where fishers are
operating have highest densities of target species and
resulting associations with environmental factors
could be used to predict fish suitable habitats (Morris
and Ball, 2006; Zwolinski et al., 2011). In the present
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
study, we further recognized inherent sources of biases
from limitations on spatial coverage of fishery data and
the MaxEnt model’s ability to predict squid potential
fishing habitat, given fishery-dependent information.
For instance, Japanese squid jigging fleets are operating
in waters west of 150°E from December–February and
across 165°E–160°W from May–July. However, Chi-
nese squid jigging fleets occupy regions across 155–
165°E from August–November, following the provi-
sions of the Sino-Japanese fishing agreement that took
 Seasonal potential fishing ground prediction of neon flying squid 199

Figure 7. The spatial distribution of squid fishing locations (colored dots) and predicted ranges of sea surface temperature (SST)
(solid lines) and sea surface salinity (SSS) (dashed lines) for January and February 2001–2004 overlain on habitat suitability
maps projected using 2001 regional models.

effect in 2000 (Chen et al., 2008). Sample selection
bias also results in environmental bias and could intro-
duce estimation errors in presence-only models includ-
ing MaxEnt (Phillips et al., 2009; Syfert et al., 2013).
Results of regional MaxEnt base models showed
minimal variability in statistical performance regard-
less of the differences in modeling domain (Table 1).
AUCs higher than 0.9 for regional base models suggest
that models have good agreement with the testing
data. AUCs from the evaluation data (Table 2) also
reinforced MaxEnt models’ good predictive ability in
inferring potential squid fishing grounds. However,
January–February (winter) fishing habitat predictions
from 2002 to 2004 showed higher AUCs than that
from June–July (summer). This difference could arise
from bias in the extent of the region sampled for base
models used in June (175–160°W; 37.5–42°N) and
July (170–160°W; 40–45°N) and wider extent of pre-
diction (150°E-160°W). Predictions of squid potential
fishing habitat from January–February (Fig. 7) also
showed better spatial matches with actual fishing
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
positions relative to June–July predictions (Fig. 8),
when the squid fishing positions used for base models
(2001) were localized to smaller regions of the basin.
In June–July prediction maps, potential squid fishing
habitats were located across 165°E–160°W. However,
potential squid fishing habitats between 150–165°E
(corresponding to the Chinese fishing grounds; Tian
et al., 2009b) were not captured as these regions were
poorly sampled in the present study and environmen-
tal conditions between 145°–165°E and of 165°E–
160°W could be quite different, possibly as a result of
the east-west gradient in Kuroshio and Oyashio Cur-
rents (Yasuda, 2003). In addition, the Chinese fishing
period extends from July to September off 145–165°E,
highlighting the spatial and temporal mismatches with
the scope of the present study. As such the summer
SST and SSS model-derived ranges between the Chi-
nese fishing ground (16.6–19.6 °C and 33.10–33.55;
Tian et al., 2009b) were different from the Japanese
fishing ground (10–17 °C and 33.5–34.3; Fig. 6)
derived from this work. In spite of data limitations and
200 I. D. Alabia et al.

Figure 8. The spatial distribution of squid fishing locations (colored dots) and predicted ranges of sea surface temperature (SST)
(solid lines) and sea surface salinity (SSS) (dashed lines) for June and July 2001–2004 overlain on habitat suitability maps pro-
jected using 2001 regional models.

Table 2. Computed area under the curve (AUC) from were sufficiently sampled in base models). MaxEnt
monthly models for January–February and June–July 2002– models, however, were unable to satisfactorily extrapo-
2004 projected using the 2001 base models. AUCs for the late the squid fishing grounds across regions with dif-
2001 base models are presented in bold. ferent environmental conditions from the base
AUC models. Thus, MaxEnt’s utility for operational fishery
applications (i.e., higher prediction accuracy) could
Model 2001 2002 2003 2004 probably be improved by circumventing data and sam-
January 0.942 0.963 0.756 0.742 pling limitations.
February 0.984 0.801 0.846 0.841 Finally, model contributions of environmental vari-
June 0.983 0.804 0.822 0.756 ables used to predict potential squid fishing habitat
July 0.969 0.793 0.589 0.847 will be discussed. In all the regional models, SST and
SSS were the most important oceanographic factors
affecting the potential fishing habitat, followed by
sampling bias in this study, MaxEnt models showed a SSH, EKE and WSC (Fig. 5). The high primary SST
strong predictive ability (i.e. based on the statistical and SSS model contributions are consistent with find-
performance (Tables 1 and 2) and modest spatial cor- ings of earlier studies showing their robust influence
respondence (Figs 7 and 8) in predicting seasonal on O. bartramii distribution and abundance in the
potential squid fishing grounds at least in sites that North Pacific (Yatsu and Watanabe, 1996; Bower and
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
 Seasonal potential fishing ground prediction of neon flying squid
Ichii, 2005). SSH and EKE data were used to deduce
the relationship between potential squid fishing habi-
tat and associated mesoscale variability (e.g. eddies
and meanders). Such oceanographic structures are
potential aggregating mechanisms for plankton and
mid-top trophic predators (Bakun, 2006; Tew Kai
et al., 2009; Mugo et al., 2014). Moderate (June;
Fig. 6) to high (January-February; Figs 5 and 6) EKE
suggests that jigging occurred in regions associated
with eddies. In the western North Pacific, eddies are
shown to influence the formation of fishing grounds
for pelagic fisheries including O. bartramii (Watanabe
et al., 2004; Zainuddin et al., 2006). Moreover, in
other ocean basins, such as Southwest Atlantic, higher
catches of the neon flying squid around the South
Atlantic Subtropical Convergence were also associ-
ated with eddies and meanders from the Brazil-Malv-
inas confluence region (Brunetti and Ivanovic, 2004).
Lastly, WSC is an indicator for surface divergence
(upwelling) and convergence (downwelling) and these
processes impact marine populations (Shanks and Eck-
ert, 2005; Morgan et al., 2009). In the present study,
MaxEnt models (Fig. 5) showed that WSC has the
least contribution to potential squid fishing grounds.
However, predominantly negative mean WSC during
winter (Fig. 3f) implies that downwelling pulses could
have taken place in squid jigging locations and could
form potential aggregation zones.
CONCLUSIONS
We have shown the feasibility of MaxEnt to elucidate
seasonal potential squid fishing grounds and eluci-
dated some possible mechanisms that could charac-
terize a potential squid fishing ground in the western
and central North Pacific. The observed aggregations
of squids at certain environmental ranges underpin
squids’ tendencies to use oceanographic features and
environmental cues that are related to presumably
high densities of forage. The effects of environmental
factors on potential squid fishing grounds also accen-
tuate changes in its habitat conditions, across and
along the zonal and meridional regions of the basin.
While this work presented a preliminary attempt to
use a machine-learning algorithm to deduce the neon
flying squid fishing grounds in the western and central
North Pacific, future comparative modeling studies
would be desirable to assess and maximize the predic-
tive capabilities of available statistical algorithms for
potential fishing habitat modeling. The inclusion of
relevant environmental factors at finer spatio-tempo-
ral resolutions and more data of squid occurrences in
the study region could also yield parsimonious
© 2015 John Wiley & Sons Ltd, Fish. Oceanogr., 24:2, 190–203.
201
forecasting models with a higher predictive power.
These insights could be useful in developing robust
potential fishing ground models for operational fishery
applications.
ACKNOWLEDGEMENTS
This work was supported by the Research on Climate
Change Adaptation (RECCA) Project of the Grant-
in-Aid from the Japan’s Ministry of Education, Cul-
ture, Sports, Science and Technology (MEXT). We
are grateful to two anonymous reviewers for their con-
structive comments on the earlier version of this man-
uscript. The authors also acknowledge the following
agencies for providing the data used in our analyses:
Aomori Prefectural Industrial Technology Research
Center for the squid fishery data; Physical Oceanogra-
phy Distributed Archive Center (PODACC) at the
Jet Propulsion Laboratory (JPL) for the AVHRR-OI
SST; NOAA Environmental Research Division’s Data
Access Program (ERDDAP) for the QuikSCAT-
derived WSC data and Archiving Validation and
Interpretation of Satellite Data (AVISO) for SSH and
geostrophic velocity data.
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