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Daylight vertical segregation of young-of-the-

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DOI: 10.1016/j.fishres.2014.03.014

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Daylight vertical segregation of young-of-the-year Argentine hake

Merluccius hubbsi: Advances in assessment of juvenile abundance
with acoustic methods
G.L. lvarez-Colombo a,b, , C.V. Dato a , L. Machinandiarena a,c ,
F. Castro-Machado a , P. Betti a
a
Instituto Nacional de Investigacin y Desarrollo Pesquero (INIDEP), Paseo Victoria Ocampo No. 1, Escollera Norte, Mar del Plata, Argentina
b
Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata (UNMdP), Funes, 3350 Mar del Plata, Argentina
c
Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires (UBA) , Intendente Giraldes 2160Ciudad Universitaria, Buenos Aires, Argentina

a r t i c l e i n f o a b s t r a c t

Article history: Assessing juvenile Argentine hake stocks from the area swept by bottom-trawls has systematically pro-
Received 5 August 2013 duced underestimates, given the adult population levels recruited from them. To gain insight into the
Received in revised form 17 March 2014 behavior and vertical distribution of rst-year Argentine hake, acoustic data and both pelagic and bottom-
Accepted 18 March 2014
trawl net samples were collected in the nursery ground of the southern stock. Five surveys were carried
Available online xxx
out in the austral winters and summers from June 2011 to February 2013. Daylight echograms at 38 kHz
showed horizontally extended aggregations of young-of-the-year (YOY) hake as layers 1020 m thick
Keywords:
separated from the seabed at average distances of 1520 m in winter and 1015 m in summer. They were
Merluccius
Young-of-the-year
above the reach of the bottom trawl nets routinely used in hake assessments. While close agreement of
Vertical distribution total length distributions was found between net samples and acoustic sh-track data in the juveniles
Acoustic assessment rst winter, only the smaller fraction of the bimodal distribution shown by the acoustic data was present
Patagonia in pelagic trawl catches six months later, in the following summer. That suggests the larger sh can avoid
the survey nets. Detection and assessment with acoustic methods allowed determination of the mean
densities and horizontal distribution patterns of early juveniles in their main nursery ground. The mean
juvenile densities obtained with acoustic methods were above one order of magnitude higher than those
obtained by bottom trawls using the swept-area method.
2014 Elsevier B.V. All rights reserved.

1. Introduction Of two main stocks, the southern or Patagonian stock is the

Hakes of the genus Merluccius constitute one of the most
intensely exploited groups of demersal sh, being primarily caught
using bottom trawls (see Lloris et al., 2005). Among them, the
Argentine hake Merluccius hubbsi supports the major demersal n-
sh shery in the Argentine Sea. Since the 1990s, more than 90% of
the western Atlantic hake catch has been comprised of this species
(Murua, 2010).
Corresponding author at: Instituto Nacional de Investigacin y Desarrollo
Pesquero (INIDEP), Paseo Victoria Ocampo No. 1, Escollera Norte, Mar del Plata,
Argentina. Tel.: +54 223 4862586; fax: +54 223 4861830.
E-mail addresses: acolombo@inidep.edu.ar (G.L. lvarez-Colombo),
cvdato@inidep.edu.ar (C.V. Dato), lauram@inidep.edu.ar (L. Machinandiarena),
fcastro@inidep.edu.ar (F. Castro-Machado), paolabetti@inidep.edu.ar (P. Betti).
most abundant, accounting for 85% of the total hake biomass in
the Argentine Sea (Aubone et al., 2000). It spawns from November
through March (late austral spring and summer) in waters off
Chubut province (Ciechomski et al., 1983; Macchi and Pjaro, 2003).
lvarez-Colombo et al. (2011) have shown how hake larvae, soon
after hatching, concentrate in a wide sector of the spawning area,
located off Isla Escondida and Camarones Bay areas. In the follow-
ing months, they are slowly distributed southwestward by bottom
currents to their nursery and settlement locations, particularly in
the San Jorge Gulf, considered to be the main hake southern stock
nursery (Fig. 1).
The San Jorge Gulf is located between 45 and 47 S cov-
ering a surface of 8000 nmi2 (square nautical miles, where
1 nmi2 = 3.434 km2 ). It is a semi-closed basin with a rather deep
connection to the open shelf (Tonini et al., 2006). Vertical mixing by
wind and tides and the entrance of southern coastal waters from the
Magellan Strait characterize the gulf, generating frontal systems.

http://dx.doi.org/10.1016/j.shres.2014.03.014
0165-7836/ 2014 Elsevier B.V. All rights reserved.

Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
G Model
FISH-3850; No. of Pages 11 ARTICLE IN PRESS
2 G.L. lvarez-Colombo et al. / Fisheries Research xxx (2014) xxxxxx
Fig. 1. Sector of the North-Patagonian shelf comprising the spawning grounds (Isla
Escondida and Camarones Bay areas) and the main nursery ground in the San Jorge
Gulf of the southern stock of Argentine hake (Merluccius hubbsi). The dashed-line
box indicates the study area selected for the distribution and acoustic numerical
density estimates of young-of-the-year hake. Dotted line indicates an example of
the acoustic track during one of the summer cruises. The bathymetry was created
by interpolation of acoustic bottom depth data (INIDEP Hydroacoustics Laboratory
bathymetry database).
Those produce nutrient enrichment and retention mechanisms
of ecological importance (Acha et al., 2004; Bogazzi et al., 2005;
Guerrero and Piola, 1997). In this productive area, the hake sh-
ery partially overlaps with that on the very important red shrimp
(Pleoticus muelleri Bate, 1888) shery. Because this sector acts as
an important hake nursery ground, high levels of juvenile hake are
discarded as incidental catch (Dato et al., 2006; Gngora et al., 2009,
2012).
As in other sectors of the Argentine Sea, hake stock assessment
in the San Jorge Gulf area is carried out by means of bottom trawl
nets hauled with the headrope a few meters above the seaoor, a
zone inhabited by most of the two-year-old and adult hake during
daylight hours (Madirolas et al., 2011). However, the vertical dis-
tribution of early juveniles in this nursery area is poorly known.
Based primarily on knowledge of the size and time of settlement
of this and other hake species, as well as from direct sampling and
otolith year-ring readings (Abella et al., 2005; Arneri and Morales-
Nin, 2000; Auster et al., 1997; Belcari et al., 2006; Buratti and Santos,
2010; Fahay, 1974; Koeller et al., 1986; Morales-Nin and Moranta,
2004; Papaconstantinou and Stergiou, 1995; Payne and Punt, 1995),
it has been assumed that young-of-the-year (YOY) Argentine hake
adopt demersal habits soon after their metamorphosis ends, mak-
ing them available to bottom nets provided with appropriate mesh
sizes.
Following these assumptions, hake juveniles including YOY sh,
have been conventionally studied from settled individuals captured
with bottom trawls during the day (Ehrlich, 1998; Ehrlich, 2000;
Ehrlich and Macchi, 2003; Ehrlich et al., 2013b; Machinandiarena
et al., 2006). However, even when bottom-trawling results have
depicted inter-annual tendencies in relative abundances and
length-class structure of age 0 and early age 1 juveniles, they
showed an evident and systematic underestimation based on the
adult population levels they had been sustaining (Castrucci et al.,
2003).
Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
Several factors may account for a downward bias of bottom-
trawl abundance estimates for young-of-the-year (YOY) juvenile
compared to the abundance levels of such juveniles required to
generate the later catches of new recruits. Those factors include (i)
net selectivity and (ii) the difference between vertical distribution
of YOY hake and the vertical aperture of the bottom sampling gear.
This paper addresses the second issue, presenting new ndings
regarding daylight behavior of YOY hake, with evidence based on
acoustic records and pelagic net tows. We provide new insight into
the behavior and vertical distribution of juvenile Argentine hake
up to one-year-old sh. The hypothesis is tested that the gradual
change from the pelagic to the demersal habitat by Argentine hake
juveniles takes longer than previously recognized for this and other
hake species. This hypothesis explains the impact of delayed set-
tlement on their catchability with bottom trawls, and it shows the
suitability of simultaneous acoustic surveys to improve assessment
of early juvenile Argentine hake abundance.
2. Materials and methods
Acoustic and biological samples were collected during ve sur-
veys carried out from June 2011 to February 2013 in the austral
winter and summer, covering the spawning and nursery grounds
(Fig. 1). The sampling happened to include the different growing
conditions experienced by the 2011 and 2012 cohorts of the south-
ern hake stock. The eld work comprised a sampling sequence
intended to evaluate methods for estimating abundance of pelagic
YOY hake. Operations included the initial characterization and
ground-truthing of the echo-recordings, delimitation of YOY hake
distribution during 2011 and the acoustic estimation of numerical
densities. With the exception of a hake prerecruit assessment sur-
vey (survey code OB0711), all the surveys considered in this study
were done in parallel with the routine bottom-trawl assessments
carried out annually during winter-spring in the Hake Global Sur-
vey and during summer, focused on the assessment of recruited
juveniles and the size of the spawning stock (Table 1).
2.1. Acoustic sampling
Acoustic data were recorded continuously between bottom
trawl stations on all cruises, sampling at ping rates variable as a
function of depth. With the exception of surveys coded EH0311 and
EH0512 (winter Global assessments) the acoustic survey design
consisted of transects regularly separated by about 20 nmi ori-
ented perpendicular to the bathymetry, encompassing a maximum
area of ca. 35,000 nmi2 comprising the spawning and nursery areas
between 43 and 47 S (Fig. 1). The acoustic equipment, calibrated
using standard sphere techniques (Foote et al., 1987), consisted
of Simrad EK500 echosounders (Kongsberg Maritime AS, Kongs-
berg, Norway) with 38 kHz (split beam, 3 dB angle of 7 , 0.3 ms
pulse duration), 120 kHz (split beam, 3 dB angle of 7 , 1 ms pulse
duration) and 200 kHz (single beam, 3 dB angle of 7 , 0.6 ms pulse
duration, not available in the R/V Capitn Oca Balda) hull-mounted
transducers. Acoustic data were logged into an INGRES relational
database during the entire surveys, and also as raw data les (sam-
ple angle and sample power) during the pelagic net operations.
2.2. Biological sampling
Pelagic tows were carried out during daylight hours with
the purpose of identifying 38-kHz sound scattering layers (SSL)
observed at Sv thresholds below 60 dB (Fig. 2). Ground truth from
the acoustic layers was obtained by means of an IKMT (Isaacs-Kidd
Midwater Trawl; 10 m2 opening) and a Nichimo-type pelagic trawl
(70 m2 opening, 22 mm mesh at the cod end).
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Table 1
Surveys analyzed, including season, acoustic and net sampling details.
Survey code Main goal Season
OB0711 Hake prerecruits Winter 2011
EH0311 Hake global assessment Winter 2011
EH0112 Hake juveniles/spawning stock Summer 2012
EH0512 Hake global assessment Winter 2012
EH0113 Hake juveniles/spawning stock Summer 2013
Survey code = ship name; survey number; year.
As a general rule, nets were placed carefully from above the
aggregations of juveniles to a maximum depth of 5 m above the
seaoor in order to avoid contamination with near-bottom sh.
The nets were trawled horizontally during 15 min at a speed of
3.5 knot (nmi h1 ) at the acoustically detected aggregation depths,
and monitored in real-time with acoustic telemetry sensors (Scan-
mar AS, sgrdstrand, Norway) to control the trawling depth and
the geometry of the net opening. All individuals caught in the IKMT
and a sample of 300500 individuals from the pelagic trawl catches
were measured to the nearest centimeter.
In addition to the ground truth pelagic trawls, catches and size
distributions of YOY hake obtained from two types of bottom trawls
during predened stations of the routine assessments, were also
considered (Fig. 2). These data allowed to compare the total length
(TL) distributions and numerical density estimates of settled pre-
recruits with those of the pelagic fraction. During survey OB0711
(winter 2011, Table 1), a small shrimp-type trawl was employed
(6 m head rope; 80 cm vertical opening; 10 mm mesh at the cod
end). This net has been utilized since 1996 to study settled pre-
recruits in the Argentine Sea (Ehrlich, 1998). During the rest of
the surveys, an Engel bottom trawl (35.3 m head rope; 5 m vertical
opening, cod end provided with 22 mm inner mesh) was operated
to assess the hake stock. Bottom nets were towed at average speeds
of 2.5 knot (shrimp-like trawl net) and 4 knot (Engel trawl) and
also monitored in real-time by means of Scanmar acoustic sensors
to control depth and the geometry of the net opening. As for the
pelagic nets, bottom sampling was performed during the day, when
the different hake size-classes adopt their diurnal behavior.
Fig. 2. Position of the pelagic trawls made to identify the composition of the acoustic
aggregations detected in the 38 kHz echograms, and the bottom trawls analyzed for
the comparison of abundance estimates.
Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
3
Acoustics Bottom sampling Pelagic sampling
Yes Shrimp-type trawl net IKMT
Yes Engel trawl net IKMT
Yes Engel trawl net none
Yes Engel trawl net Pelagic trawl net
Yes Engel trawl net Pelagic trawl net
2.3. Echogram analysis and echo-integration measurements
Based on the characterization of the pelagic aggregations iden-
tied as YOY hake, their contribution to total 38 kHz backscattering
was estimated and recorded during the analyzed surveys. This
interpretation of the echograms using BEI software (Bergen Echo-
Integrator; Korneliussen, 2004) involved analysis of the water
column into pelagic, surface-referenced channels and a bottom-
referenced channel comprising the deepest 10 m of the water
column, above the acoustic dead-zone (ADZ; Ona and Mitson,
1996). Echo-integration measurements in terms of the nautical
area scattering coefcient (NASC, in m2 nmi2 ; MacLennan et al.,
2002) at a volume backscattering strength (Sv ) threshold of 80 dB,
were averaged over elementary sampling distance units (ESDU)
of 1 nmi (nautical mile = 1.852 km) and exported for spreadsheet
analysis in 5 m, surface-referenced channels. The contribution of
stronger acoustic targets, mainly adult hake, was measured at an Sv
threshold of 60 dB and subtracted. Only daylight echograms were
considered for this study, given the well-known DVM behavior of
advanced juveniles and adults that are strongly demersal during
daylight (see Ehrlich, 1998; Ehrlich et al., 2013a).
The NASC values integrated in the 5 m, surface-referenced chan-
nels were converted to bottom-referenced data and employed to
determine the average distance from the seabed of the pelagic
aggregations. Given that YOY juveniles in the deepest 5 m channel
could be acoustically hidden by the adult and settled-juvenile frac-
tions, and also by the ADZ effect, their acoustic contribution in that
layer was estimated from the bottom catches with the Engel net.
To estimate the contribution of each 1 cm size-class (i subscripts)
of juveniles <20 cm to the total NASC in the deepest 5-m layer, their
absolute frequencies in the catches (fi ) were calculated following
Bodholt (1990). Thus, the areal density of each size-class (i ) was
estimated considering its percent abundance in the catches and the
corresponding theoretical backscattering cross-section (bsi ) as
fi
i = n NASC (1)
f
i=1 i
bsi
NASCi = i bsi (2)
where bsi was obtained from the target strength to length (TS/TL)
relationship established by Madirolas and Castro-Machado (2008)
for this species:
TS (dB) = 39.3 log TL (cm) 97.1 dB (3)
Finally, the equivalent NASC fraction of the juveniles <20 cm was
estimated by summing of the resultant NASCi values.
2.4. Acoustic single-target analysis
In order to estimate the numerical abundance of YOY hake
in the pelagic aggregations and to study the relation of sh size
with depth, the acoustic target-strength (TS) from individual shes
detected in the aggregations were examined. The raw data les
recorded during the operation of the IKMT and the pelagic trawl net
were analyzed by means of Miryax Echoview v4.1 (Myriax Pty Ltd.,
Hobart Tasmania, Australia) post-processing software. By means
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4 G.L. lvarez-Colombo et al. / Fisheries Research xxx (2014) xxxxxx
Table 2
Echoview software settings employed for single target and sh-track analysis.
Single target detection settings
TS threshold (dB) 80
Pulse length determination level (dB) 3.0
Minimum normalized pulse length 0.4
Maximum normalized pulse length 1.5
Maximum beam compensation (dB) 12
Maximum standard deviation of angles (degrees) 0.1
Filter angles for single targets (degrees) 3 Finally, mean TS of the sh-tracks detected during winter and
Fish tracking settings
Algorithm data Range, angles and time
Track detection (alpha) 0.7
Track detection (beta) 0.5
Target exclusion distance (m) 4 mated for the settled YOY juveniles caught with bottom trawls in
Weight (major and minor axis) 30
Weight (range) 40
Minimum number of single targets in a track 4
Minimum number of pings in a track 4 3. Results
Maximum gap between single targets (pings) 0
TS = target strength; dB = decibel.
of the Virtual Echogram and Fish Tracking modules of the soft-
ware (parameter settings detailed in Table 2), the echo-signatures
from individual shes were isolated, and the individual TS values
averaged.
The data analysis included procedures intended to minimize
two different sources of overestimation of the TS estimates. Thus,
a cut-off angle of 3 was set to the single target detection algo-
rithm given that the signal-to-noise ratio is progressively reduced
with increasing angle from the acoustic axis, and hence the esti-
mates of TS are positively biased outwards it (Fleischman and
Burwen, 2000). Second, due to the increasing tendency towards
inclusion of multiple targets with increasing range from the trans-
ducer, the Fish Tracking algorithm was conservatively set, allowing
only sh-tracks composed of at least four consecutive single-target
detections (Table 2). Finally, considering that only one target can
occupy the acoustic resolution volume for the measurement to
be valid, the relation between NASC, range of the identied sh-
tracks and TS was employed to check that multiple targets were not
present in the detection volume, setting an upper limit on operation
of the single target detector, following Ona and Barange (1999):
NASC(c/2)r 2 D
N=
410TS/10 z(1852)2
where c is the sound speed;  is the pulse duration; r is the range;
z is the depth interval; and D = 2(1 cos  D ) is the sampled
volume for a solid angle of  D = 3 (Table 2).
The mean TS of the pooled targets detected in the layer 530 m
off the bottom was employed in abundance estimations, whereas
the average TS of individual sh tracks were converted to sh
length, following the relationship established in Eq. (3), in order
to study a possible relation between sh total length and depth.
2.5. Horizontal distribution patterns and numerical density
estimates of YOY hake
The NASC measurements were integrated over the water-
column, interpolated over the sampling track and mapped. Given
the visual evidence of spatial structure in the horizontal distribu-
tion of hake pelagic juveniles over the study area, geostatistical
analysis was employed to analyze sh-echo spatial autocorrela-
tion and to evaluate the variance of the abundance estimates. An
exponential model was tted to the observational variograms of
the distance vector (h). The model had a practical range equal
to three times its range parameter (a) and employed maximum
Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
likelihood estimations for the variogram parameters sill (C) and
range (a) (Rivoirard et al., 2000). The specic model was tted as
 
|h|
(h) = C[1 exp a ]. (5)
EVA v2.0 software (Estimation VAriance; Petitgas and Prampart,
1993) was employed to estimate the geostatistical variance of the
mean NASC variable.
summer 20112013 were employed together with the mean echo-
integration value of each survey to estimate the numerical densities
of pelagic YOY juveniles for comparison with the densities esti-
the San Jorge Gulf.
3.1. Echogram characterization and delimitation of the
distribution area
Daylight echograms obtained at 38 kHz during the rst survey
showed horizontally extensive aggregations of scatterers in layers
1020 m thick mainly conned to the nursery ground of the San
Jorge Gulf area (Fig. 3). For this reason, the comparative analysis of
the analyzed survey series is related to the Gulf area.
Even when the aggregations were occasionally observed in
proximity to the seaoor, they were usually noticeably separated
from it at distances above the reach of the bottom trawl nets rou-
tinely used in hake assessments. The vertical distribution changed
at dusk when the aggregations rose towards the surface dispersing
in the upper layers.
3.2. Bottom trawl catches
The use of the shrimp-type bottom trawl during the rst winter
survey (OB0711), showed that settled YOY juveniles were present
in most of the study area with a numerical density average of
36,454 ind nmi2 (SD = 53,829; n = 105). The uniform coverage of
the study area with this net allowed for a comparison between
the horizontal distributions of settled YOY juveniles with that
of the pelagic fraction, obtained from the acoustic data. While
the highest density values of settled individuals (maximum of
(4)
253,364 ind nmi2 ) was located outside the Gulf, the acoustic
detected pelagic aggregations were mostly recorded inside the Gulf
(Fig. 4).
Length distributions derived from the Engel trawl in San Jorge
Gulf catches, showed that both in winter 2012 and summer 2013
less than 10% of total numerical abundance corresponded to YOY
juveniles 15 cm TL.
3.3. Pelagic trawl catches
Pelagic trawls were restricted to the San Jorge Gulf, where most
of the acoustic records related to YOY hake were registered.
Maximum catches of YOY hake obtained in all of the IKMT
deployments in winter 2011 were 100 juveniles in 15-m tows.
However, strong avoidance of the net should be expected, consid-
ering the rather small sampler mouth, the bridles in front of it
during operation and the fact that shing was during daylight. Com-
parison of the IKMT and the shrimp-type bottom trawl catches in
the Gulf showed similar TL distributions of YOY juveniles (Fig. 4),
with means of 6.78 cm (SD = 2.98; n = 385) and 7.15 cm (SD = 2.57;
n = 448), respectively. However, a statistically signicant difference
between group means was determined by one-way ANOVA (F(1,
831) = 10.13, p = 0.0015).
G Model
FISH-3850; No. of Pages 11 ARTICLE IN PRESS
G.L. lvarez-Colombo et al. / Fisheries Research xxx (2014) xxxxxx 5

Fig. 3. Example of a daylight echogram recorded in San Jorge Gulf (summer 2012). The arrows indicate an aggregation of young-of-the-year (YOY) hake (Merluccius hubbsi)
at a mean distance of 25 m off the seaoor and sh-tracks of adult hake in the bottom channel (time labels in bottom channel regions in UTC-3).

Large catches, almost exclusively composed of hake YOY
juveniles were obtained with the pelagic net in 20122013, con-
rming the composition of the studied aggregations. The by-catch
consisted of a few individuals of other species like jellysh (Mne-
miopsis leidyi; Aequorea sp.), squid (Loligo gahi), sh (Stromateus
brasiliensis; Engraulis anchoita) and crustaceans (Munida gregaria
and Peisos petrunkevitchi, related to acoustic scattering layers
mostly recorded at the higher frequencies).
In winter 2012, an average of 1600 sh was caught in the
15-m tows of the pelagic net (SD = 947; n = 8). Distribution of
the pooled TL data was unimodal with a mean TL of 8.01 cm
(SD = 1.48; n = 13,059). In summer 2013, an average of 1300 pelagic
juveniles were caught in similar 15-m tows (SD= 473; n= 5),
also showing a unimodal distribution of TL averaging 11.29 cm
(SD = 1.96; n = 6574) (Fig. 5). Comparison between winter and
summer TL means showed signicant differences (ANOVA (F(1,
13168) = 11655, p < 0.0001) as expected considering the juvenile
growth during the ve months elapsed between surveys.
3.4. Acoustic sh-tracking
The Fish-track module of Echoview software allowed the detec-
tion of single targets that could be identied as YOY hake using
data from the depth-directed sampling with the pelagic trawl.
In winter 2012, a unimodal distribution was obtained from the
pooled data with modal TS at 58 dB, while in summer 2013
a bimodal distribution with modal TS at 56.5 dB and 46.5 dB
was found. The conversion of acoustic sh-track TS data to sh
length, employing the empirical TS/TL model, showed close dis-
tributions of acoustic and directly measured TL in winter, even
though with a statistically signicant difference between group
means (ANOVA (F(1, 26118) = 5661, p < 0.0001). In summer, the TL
distribution of the sh caught with the pelagic trawl only accounted
for the lower TL mode of the distribution of TS/TL converted values
(Fig. 5).
3.5. Vertical distribution
The average vertical distribution of NASC values in the 5 m
bottom-referenced channels revealed YOY hake aggregations
located above the seaoor at variable distances. In winter a NASC
peak was found at 1520 m, with some echo traces reaching as
much as 50 m off the bottom. In summer the aggregations were
closer to the seaoor with their NASC peak at 1015 m above it
(Fig. 6).
The mean depth of the tracks for YOY hake of different sizes
(acoustic signals converted to length from the TS to TL relation-
ship), shows a progressive shift toward the bottom with increasing
TL, as exemplied in Fig. 7 for winter and summer (surveys OB0711
and EH0113, respectively). This nding resembles the differential
seasonal depth distribution shown in Fig. 6 and supports a grad-
ual, downward ontogenetic shift in daylight location during the
settlement process.
3.6. Horizontal distribution patterns and abundance estimates of
YOY hake
The acoustic data interpolated over the study area showed that
NASC was not distributed randomly but exhibited well-dened pat-
terns. That is shown in the empirical variograms for winter and
summer cruises, as well as in the map plots which exhibit distinc-
tive areas of higher concentration in the northern and southern
sections of the San Jorge Gulf (Figs. 8 and 9; Table 3). The vari-
ograms show an autocorrelation range around 2555 nmi in the
along-transect direction, giving a preliminary indication that the
distributions of the acoustic sampling tracks were adequate to
reveal the underlying spatial structure.
The density estimates for YOY hake in winter (2011 and
2012) and summer (2012 and 2013) had similar seasonal aver-
ages all across the San Jorge Gulf region, with approximately
1200,000 ind nmi2 in winter and 300,000 ind nmi2 in summer
(Table 4). The smaller mean density values obtained for the pelagic

Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
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Fig. 4. Results obtained in the rst survey (June 2011). Above left: horizontal distribution of settled young-of-the-year (YOY) hake (Merluccius hubbsi) with density scale
in ind nmi2 . Dots indicate the geographical position of the shrimp-type trawl sampling stations). Below left: Total length (TL) distribution obtained from the bottom net
catches. Above right: numerical densities obtained from the echo-integrated 38 kHz acoustic signals with density scale in ind nmi2 . Below right: Total length (TL) distribution
obtained from the Isaacs-Kidd Mid-water Trawl (IKMT) catches.

Table 3 aggregations in summer reect both the advance of the settlement

Statistical parameters of the exponential model tted to the experimental
process and the greater juvenile TL with correspondingly greater
variograms.
TS values (Table 4).
Winter Winter Summer Summer The mean densities obtained with acoustic methods were com-
2011 2012 2102 2013
pared with those obtained by the swept-area method. A mean
Sill (C) 2802.32 1418.20 1482.95 1571.92 density of 17,508 ind nmi2 was obtained with the shrimp-type
CI (95%) low 2661.13 1353.72 1272.72 1432.07 trawl catches in the Gulf area in 2011. That is nearly two orders
CI (95%) upp 2943.50 1482.69 1693.18 1711.77
of magnitude less than the concurrent acoustic estimate. Similarly,
CV (%) 2.19 1.98 6.16 3.87
Range parameter (a) 10.41 8.16 11.24 18.70 the density estimates for juveniles <15 cm obtained with the Engel
CI (95%) low 8.29 6.40 5.13 14.25 trawl were very much lower than indicated by the acoustic tech-
CI (95%) upp 12.53 9.92 17.36 23.14 niques.
CV (%) 8.85 9.35 23.64 10.34
Range (3a) 31.22 24.48 33.73 56.09
CI (95%) low 24.86 19.21 15.38 42.75 4. Discussion
CI (95%) upp 37.58 29.75 52.07 69.43
R2 0.84 0.77 0.41 0.84
Previously, Argentine hake have been considered to share com-
VAR 29.50 18.53 19.86 15.20
mon patterns of development and habitat use with other hake
CI (95%) low = lower condence limit at the 95% level; CI (95%) upp = upper con- species, in particular the early stage and small size at settlement.
dence limit at the 95% level; CV = coefcient of variation; R2 = coefcient of
determination; VAR = geostatistical variance estimate.
Our data show, however, that a high portion of the early juveniles of
the Argentine hake southern stock aggregate during daylight well

Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
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Fig. 5. Average total length (TL) distributions of hake (Merluccius hubbsi) early juveniles obtained in winter 2012 and summer 2013 with the pelagic trawl net (above) and
by the conversion of acoustic sh-tracks mean target strength to TL (below).

above the seaoor during a protracted period during their rst year
of life. This is likely the main factor affecting the apparent under-
representation of this size fraction in demersal trawl catches during
swept-area assessments. This behavioral feature also implies the
likelihood that they can be detected and assessed with acoustic
methods. The particular characteristics of the north Patagonia sec-
tor of the Argentine Sea, with its low sh diversity and the relatively
high abundance of the Argentine hake, allowed us to study these
aspects of its life history with a straightforward approach.
Several biotic and abiotic factors, such as the predator-prey
interactions of YOY hake and the circulation features in the study
area, may account for the observed daylight distribution. It has been
demonstrated for other hake species that, after an early settlement,
YOY juveniles perform diel vertical migrations (DVM) from near-
bottom depth towards the surface during darkness following their Fig. 6. Average nautical area scattering coefcient (NASC) distributions of bottom-
prey, mainly planktonic crustaceans like euphausiids (Bowman and referenced, 5-m echo-integration channels corresponding to hake (Merluccius
Bowman, 1980; Bozzano et al., 2005; Koeller et al., 1989; Pillar and hubbsi) early juveniles in winter 2012 and summer 2013. The dotted line indi-
Barange, 1997). Juvenile M. hubbsi in the nursery ground of San cates the maximum vertical aperture of the Engels bottom trawl net employed for
Argentine hake stock assessments.
Jos Gulf mainly feed on the amphipod Themisto gaudichaudi and
the euphausiid Euphausia lucens (Snchez and Garca De la Rosa,
1999; Temperoni et al., 2013). Layers and swarms of these zoo- Avoidance of predation from settled conspecics is another fac-
plankters are commonly observed at higher acoustic frequencies tor governing the vertical segregation of YOY hake in the nursery
(120200 kHz) to inhabit intermediate depths during the day, often ground. The cannibalistic behavior of gadoids in general and hakes
in close contact with pelagic hake juveniles. Thus, the hake YOY in particular has been extensively treated in the literature (see Link
juveniles are able to feed during daytime. Ongoing studies of the et al., 2012 and references therein). While the diet of juvenile hake
stomach contents of juveniles from our pelagic catches will provide is primarily small crustaceans, larger hake become piscivorous,
new information on the feeding strategy of YOY hake as a function with cannibalism often becoming the dominant source of predation
of their vertical distribution pattern. mortality on YOY classes, with potential impacts on recruitment

Table 4
Numerical density estimates of young-of-the-year (YOY) hake in winter and summer, and the acoustic parameters nautical area scattering coefcient (NASC) and target
strength (TS) employed.

Survey Fish-tracks mean TS (dB re 1 m2 ) Mean  bs (m2 ) N SD Mean NASC (m2 nmi2 ) Mean density (ind nmi2 )

Winter 2011 55.10 3.09E 06 12858 3.55 43.33 1116,196

Winter 2012 56.38 2.30E E 06 6806 3.25 33.22 1149,266
Summer 2012 53.46 4.51E E 06 2157 3.77 17.53 309,468
Summer 2103 50.52 8.88E E 06 2207 4.89 28.86 258,758

TS = target strength; dB = decibel;  bs = acoustic backscattering cross section; n = number of sh tracks analized; SD = standard deviation; nmi = nautical mile.

Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
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8 G.L. lvarez-Colombo et al. / Fisheries Research xxx (2014) xxxxxx
Fig. 7. Average off-bottom distances and 95% condence intervals of the sh-tracks
assigned to hake (Merluccius hubbsi) juveniles, detected between 5 and 30 m above
the bottom in winter 2012 (above) and summer 2013 (below).
and population regulation (Juanes, 2003; Tsou and Collie, 2001;
Waldron et al., 1989). Macpherson and Gordoa (1994) found that
cannibalism in Cape hake (Merluccius capensis) is not related to prey
density but to a marked preference for small conspecics. As for the
Argentine hake it is well recognized that cannibalism is an impor-
tant part of natural mortality (Ocampo Reinaldo et al., 2011; Prenski
and Angelescu, 1993; Snchez and Garca De la Rosa, 1999). Snchez
and Garca De la Rosa (1999) showed a gradual size-related inter-
cohort cannibalism in Argentine hake with YOY juveniles being
mainly preyed upon by age 1 and age 2 settled juveniles. In this
sense, preliminary observations during the surveys analyzed in this
study show that, in areas where YOY acoustic layers approached the
bottom, the Engel trawl catches included both more small juveniles
and small juveniles in the stomach contents of larger juveniles.
The acoustic techniques employed here allowed a uniquely
detailed analysis of the relationship between YOY hake size and
depth. The relationship between the mean TS of sh-tracks and
depth implies a gradual process of settlement; the smaller sh
showed the highest average distance from the bottom. Possibly
that indicates behavior selected to minimize predation by older
juveniles, cannibalism demonstrated from the Engel trawl catches.
Various aspects of the observations conicted somewhat with
this hypothesis. Even when a cautious approach was applied for
sh-track selection, also considering the acoustic resolution vol-
ume, the positive bias from multiple scatterers being misidentied
as single sh in those sections with the highest NASC values has to
be considered as a possible source of error. Also, catches with the
bottom trawls demonstrate an early settlement of at least part of
the new cohorts.
In the example presented in this study, the TL distributions
of pelagic and settled YOY were similar, and the sector with the
highest densities of settled YOY juveniles was different from the
Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
sector with maximum pelagic concentrations (Fig. 4). Given the
batch-spawning nature of hake and the extent and variability of
the spawning area (Macchi et al., 2005, 2007, 2010), successive
broods from different times or places during the breeding sea-
son would face different environmental conditions. Some of those
would coincide with favorable conditions for development of the
early life stages, and some would not. Thus, variable support for
development could be the main cause of the observed variability in
settlement timing. Also, as has been established for many gadoid
species, including silver hake (Merluccius bilinearis), YOY juveniles
settle and concentrate over particular substrates that provide phys-
ical and visual shelter from predators (Auster et al., 1991; Auster
et al., 1997; Steves et al., 1999). In the case of settled M. hubbsi pre-
recruits, preference for certain bottom types has been observed. In
particular, seaoor characterized by high content of ne granulo-
metric fractions and silt, as well as beds of the Patagonian scallop
Zygochlamys patagonica, have been related to the highest catches of
settled YOY juveniles obtained with the shrimp-type bottom trawl
in earlier assessments (Ehrlich et al., 2001).
The association between the circulation patterns in the San Jorge
Gulf area and the vertical distribution patterns of pelagic YOY hake
remains a subject for future research. The circulation patterns of
the San Jorge Gulf, described by Tonini et al. (2006), are complex in
both the horizontal and vertical planes, which possibly accounts for
the spatial structure in the horizontal distribution of the integrated
acoustic echoes. Responding to the predominantly west wind, the
circulation consists of two opposed gyres: clockwise in the north-
ern sector and counter-clockwise and more intense in the southern
sector. In the vertical domain, there is a two-layer circulation; the
deeper layers ow toward the coast, the surface layers ow away
from it. Active positioning by larval stages in a countercurrent circu-
lation system has been postulated for the Argentine hake spawning
grounds (lvarez-Colombo et al., 2011), and it is possible that anal-
ogous positioning is adopted by the juvenile stages.
Our ndings have a direct impact on the assessment of new
cohorts of the Argentine hake southern stock, revealing the value
of acoustic techniques for survey of the nursery areas and for esti-
mating the total abundances of early hake juveniles. For other
gadoids like cod and haddock, given their wide vertical distri-
butions that often extend far above the bottom, swept-area and
acoustic indices of abundance covering different fractions of the
stock, are already being combined into single estimates of absolute
abundance, improving the reliability of survey results and provid-
ing a wider viewing frame for understanding sources of variability
(God and Wespestad, 1993; Hjellvik et al., 2003; Jakobsen et al.,
1997). This also applies to several hake stocks that have been the
subjects of combined surveys (Fleischer et al., 2005; Guttormsen
et al., 2003; Jakobsen et al., 1997). The results of this study show
that the particular behavior of the bulk of YOY hake makes them
readily available to echo-sounder detection during the day. It is
fully feasible to perform a simultaneous acoustic surveying along
with bottom trawl sampling, thus partly overcoming the time con-
straints of trawl assessment surveys across the long and wide
Argentine shelf.
Estimates of the numerical abundance of pelagic aggregations
with acoustic techniques were more than one order of magnitude
greater than those obtained with bottom trawls. However, while
in the winter surveys the TL distribution estimated for YOY hake
from the pelagic net catches resembled the unimodal distribution
based on the acoustic sh-tracks, in summer the TL distribution
based on trawl catches was skewed toward the smaller fraction of
the bimodal distribution based on acoustic data. That indicates pos-
sible avoidance of the net by the larger juveniles. Valdemarsen and
Misund (1995) analyzed the performance of different pelagic trawl
nets, concluding that a net of 10 10 m, similar to the net dimen-
sions employed in this study, is appropriate for pelagic gadoid
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Fig. 8. Left: acoustic transects (dotted lines) and interpolated horizontal distribution of the nautical area scattering coefcient (NASC). Right: experimental (dots) and tted
(line) variograms obtained in winter 2011 (above) and 2012 (below).

Fig. 9. Left: acoustic transects (dotted lines) and interpolated horizontal distribution of the Nautical Area Scattering Coefcient (NASC). Right: experimental (dots) and tted
(line) variograms obtained in summer 2012 (above) and 2013 (below).

Please cite this article in press as: lvarez-Colombo, G.L., et al., Daylight vertical segregation of young-of-the-year
Argentine hake Merluccius hubbsi: Advances in assessment of juvenile abundance with acoustic methods. Fish. Res. (2014),
http://dx.doi.org/10.1016/j.shres.2014.03.014
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10 G.L. lvarez-Colombo et al. / Fisheries Research xxx (2014) xxxxxx
juveniles of 10 cm or less. They also concluded that a larger trawl
opening is needed to capture larger sh efciently. Thus, further
trials with nets of larger dimensions and/or faster towing speeds
are needed to improve the validation of the acoustic data during
summer.
The vertical herding effect of trawling on gadoids, the result of
their strong tendency to dive just after the passage of a vessel and/or
trawl warps, has been extensively studied (Aglen, 1996; Aglen et al.,
1999; Handegard and Tjstheim, 2005; Handegard et al., 2003;
Nunnallee, 1991; Ona and God, 1990). Thus, the so-called effec-
tive shing height (EFH) (Aglen, 1996; Hjellvik et al., 2003; Kotwicki
et al., 2013), is the level at which a near-bottom trawl will catch the
most sh among those placed well above the bottom, sh observed
with hull-mounted transducers well ahead of the net being towed
behind. This behavior and the associated effect on bottom trawl
catches have been demonstrated for Argentine hake by Casarsa
and Madirolas (2007). However, the systematic underestimation of
early juveniles in the Engel net catches during assessment surveys
of the southern Argentine hake stock provides evidence of a negligi-
ble effect of vertical herding for YOY hake in pelagic aggregations, at
least during their rst year of life. This conclusion is consistent with
earlier investigations, that the vertical herding is least for small sh
(Aglen, 1996; Aglen et al., 1999). This could also partially explain
the size structure of the by-catch in commercial catches. According
to Dato et al. (2006), about 87% of Argentine hake discards, ranging
from 28 to 32% of the total number of individuals caught, were in
the Age-1 fraction. Young-of-the-year juveniles (Age-0) accounted
for less than 1%.
Acknowledgments
Special thanks to Adrin Madirolas for his valuable advice and
ideas, as well as for his contribution in the acoustic data analysis.
To Hermes Mianzan for his advice during the manuscript writ-
ing process, to Anbal Aubone and Griselda Garaffo for guidance
on the statistical analysis, and to Ral Bertelo for his commitment
to set up the pelagic nets. Success at the eldwork stage would
not have been possible without the enthusiastic support of the
crews of the INIDEP research vessels, particularly during the pelagic
net operations aboard the R/V Dr. Eduardo L. Holmberg. This work
was partially supported by the Research Project PIP CONICET 112-
200801-00815 directed by Dr. Gustavo Macchi (CONICET/INIDEP).
This is INIDEP contribution no. 1865.
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