FISHERIES OCEANOGRAPHY
10:2, 193±206, 2001
Relationship between spatial distribution of the Patagonian
stock of Argentine anchovy, Engraulis anchoita, and
sea temperatures during late spring to early summer
J. E. HANSEN,1,* P. MARTOS1,2 AND
A. MADIROLAS1
1
Instituto Nacional de InvestigacioÂn y Desarrollo Pesquero
(INIDEP), P.V.Ocampo 1, 7600 Mar del Plata, Argentina
2
Universidad Nacional de Mar del Plata, Funes 3350, 7600
Mar del Plata, Argentina
ABSTRACT
The relative abundance of adult Argentine anchovies
(Engraulis anchoita) and the thermal structure of the
sea between 41° and 45°30¢S during four research
surveys were compared. Acoustic data were collected
while making parallel transects along which CTD
2 (conductivity, temperature, depth) stations were
regularly distributed. Anchovy abundance was related
to both the sea surface temperature and the strati®-
cation of the water column, as classi®ed according
to the / parameter of stability (Simpson, 1981).
Regarding the whole water column, the sea tempera-
tures where adult anchovies were recorded ranged
from 8.5 to 16.5°C, but anchovy echo traces from
waters above 14°C were obtained mainly at night. The
sea surface temperature bounds for anchovy distribu-
tion were 11 and 17°C, with a preferential range
between 12.5 and 16°C, but no absolute value of sea
surface temperature was found to be most favourable
for the species. The highest ®sh abundances were
related to sharp thermal gradients, either horizontally
recorded in frontal zones (» 0.02°C or more per km) or
vertically associated to the occurrence of a thermo-
cline (stability > 10 J m±3). Sea fronts seemed to be
the axes for the distribution of anchovy shoals, and the
annual changes in the positions of the fronts seemed to
be followed by analogous changes in the locations of
the main anchovy concentrations. Within thermally
strati®ed areas and during the daytime, anchovies
apparently preferred the thermocline layer or the layer
*Correspondence. e-mail: jhansen@inidep.edu.ar
1 Received 4 October 1999
Revised version accepted 7 September 2000
Ó 2001 Blackwell Science Ltd.
Fish. Oceanogr.
immediately above that, although a few ®sh shoals
were located below the thermocline.
Key words: Argentine anchovy, distribution, Engraulis
anchoita, Patagonian stock, sea temperatures
INTRODUCTION
Anchovies (Engraulis anchoita) are the most abundant
and most ecologically important pelagic ®sh resources
off Argentina. Two anchovy populations are recog-
3 nized south of 34°S, the Bonaerensis and the Patago-
nian stocks (Brandhorst et al., 1974; Hansen et al.,
1984; SaÂnchez and Martos, 1989). These ®sh tolerate
wide ranges of both temperature (8±23°C) and salinity
(14±35 p.s.u.), like other species of the genus Engraulis
(Reid, 1966), but their main concentrations would
occur where those environmental variables show sharp
gradients (Hansen et al., 1986; Hansen and Madirolas,
1996). In the daytime, these ®sh (like other small
pelagic species) respond to the light by forming com-
pact shoals at different depths. On the other hand,
they are widely scattered at night, in water layers close
to the sea surface (Gudmundsson and Gamberale,
1972; Hansen and Madirolas, 1996; Castello, 1997).
The distribution area of the southern (Patagonian)
stock extends from 41 to 48°S, and mainly between
41 and 45°S during the last quarter of the year (Fig. 1).
From the oceanographic point of view, a large portion of
that zone, especially the area which is close to the
ValdeÂs Peninsula (42°30¢S), offers several conditions
that favour the formation of sea fronts in spring and
summer. Among such conditions are the occurrence of a
clear thermocline during spring and summertime
(Guerrero and Piola, 1997; P. Martos and R. A. Guer-
4 rero, unpublished data), as well as a great energy dissi-
pation due to tides (Miller, 1966; Glorioso and Simpson,
1994). The sea fronts represent the border between
strati®ed waters and those intensely mixed by tides.
Martos and SaÂnchez (1997) have preliminarily
described three frontal zones of a seasonal nature
within the region: a thermohaline sea front located off
193
 194 J.E. Hansen et al.

Figure 1. Distribution area and survey design for the study of the Patagonian stock of Argentine anchovy during late spring to
early summer. Grey lines, Strata 1±5 used to estimate anchovy biomass; dotted lines, acoustic transects; A and B, oceanographic
sections mentioned throughout the text.

the mouth of San MatõÂas Gulf (41±42°S); a tidal
front close to the ValdeÂs Peninsula; and another tidal
front that extends along the Patagonian coast, close to
Escondida Island (43°40¢S).
Sea fronts usually have great biological importance
5 because they are suitable for ®sh spawning, as well as
for egg and larvae development. They are highly
productive zones and at times correspond to areas of
accumulation of the early stages of some ®sh species, as
they can represent geographically stable larval
Ó 2001 Blackwell Science Ltd., Fish. Oceanogr., 10:2, 193±206.
retention areas (Iles and Sinclair, 1982). It has been
shown that the spawning areas of the Patagonian stock
of Argentine anchovy are related to the above-
mentioned sea frontal systems (SaÂnchez and
Ciechomski, 1995; Martos and SaÂnchez, 1997).
Although the distribution of small pelagic ®sh is
generally associated with the main physical features of
the marine environment (Castillo et al., 1996), most
studies about adult Argentine anchovy concern the
spawning grounds. Most of the research work has dealt

with the Bonaerensis stock, which also inhabits the
southern shelf off Brazil between 28 and 34°S during
winter and springtime (Castello, 1997). The present
paper is directed to the analysis of the relationship
between the distribution of adult Patagonian ancho-
vies and thermal structure of the sea during late spring
and early summer.
MATERIALS AND METHODS
Design of the surveys
The basic data were recorded during four research
surveys directed to evaluate the Patagonian anchovy
biomass between 41° and 45°30¢S, from the coast to,
approximately, the 100 m isobath (» 42 000 nm2),
during the season that is known to be the peak of
the spawning activity of the stock (SaÂnchez and
Ciechomski, 1995). These cruises were carried out by
7 the Research Vessel `CapitaÂn Oca Balda', a 65-m-long
stern trawler, in December, every year from 1993 to
1996. All those surveys kept the design adopted in
1993, which has been already described (Hansen and
Madirolas, 1996). The area of the surveys, which
covered most of the area of anchovy distribution during
the southern late spring, was split into ®ve strata
according to previous records of relative abundance of
both egg and adult ®sh of the species. The design of the
acoustic surveys included 23 parallel transects, which
were randomly distributed into the different strata
based upon the expected variability of the above-
mentioned parameters (Fig. 1). Stratum 1 is located at
the mouth of San MatõÂas Gulf and is characterized by
the presence of a thermohaline sea front. Stratum 2 is a
large zone mostly in front of the ValdeÂs Peninsula and
southwards, which is dominated by the occurrence of a
tidal sea front. Stratum 3 comprises deep, completely
strati®ed waters. Stratum 4 includes the main spawning
ground of the anchovy stock during this season (SaÂn-
chez and Ciechomski, 1995), which is centred in
Escondida Island and related to the other coastal, tidal
sea front mentioned in the Introduction section. This
front also extends over the inshore part of Stratum 5,
which is the southernmost extreme of the anchovy
distribution in this season.
Acoustic and biological data
The acoustic equipment used in 1993 was a scienti®c
echosounder SIMRAD EK400 connected to a digital
echointegrator SIMRAD QD. The echointegrator
output was recorded in real time by using a personal
computer and the software HYDRO to store and process
the acoustic data (Madirolas and Rodriguez, 1996).
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Distribution of Argentine anchovy and sea temperature 195
During the 1994 survey, a scienti®c echosounder SIM-
RAD EK500 was used, but the original system was used
for storing and processing data. Since 1995, a Graphic
Workstation HP9000 and the postprocessing software
SIMRAD BI500 were linked to the EK500.
Hull-mounted 38-kHz transducers and a pulse dura-
tion of 1 ms were always used. To enable comparison of
the data from different years, the scienti®c sounder was
calibrated before every cruise, using a standard target
and the procedure suggested by Foote et al. (1987). The
output signal of the echosounder was processed by
echointegration (Forbes and Nakken, 1972) and echo-
integration layers of 10 m depth were de®ned, the ®rst
starting at 2 m from the transducer (i.e. 5.5 m from the
sea surface). The symbols in the ®gures displaying
anchovy distributions correspond to the square root of
the column sound backscattering coef®cient, sa (Foote
8 et al., 1991), normalized for a square nautical mile and
measured in units of m2/nm2. For every survey, anchovy
biomass estimates were derived from the values of the
column scattering coef®cient. The formulae utilized for
converting the sa values to biomass density by using the
size±target-strength relationship corresponding to
clupeoid ®sh, as derived by Foote (1987), are described
in Hansen and Madirolas (1996).
At any time of day or night during the acoustic
tracking, ®shing hauls were performed to identify echo
traces when they indicated great quantities of anchovy
or other species. A mid-water trawl net having a 10-mm
mesh size at the inner cover of the codend was used.
From 12 to 20 hauls per cruise were made. In every haul
where anchovy was caught, samples of the species were
taken (m ³ 120 individuals per haul). Besides other
biological data, total length and total weight (precision
5 mm and 0.1 g, respectively) were recorded.
Oceanographic data
Between 54 and 94 oceanographic stations were
regularly distributed in each cruise on the acoustic
tracks. Continuous pro®les of both temperature and
conductivity were obtained by either a compact CTD
9 (conductivity, temperature, depth) `Meerestechnik
Elektronik' (1993 survey) or a `Sea Bird 19' CTD
(cruises since 1994). The data series were then proc-
essed using standard Seasoft routines (Sea-Bird Elec-
tronics, 1997) and reduced to values of temperature
10 and salinity corresponding to a 1-db interval. The
CTD was calibrated by measuring water samples with a
salinometer and reversing thermometers.
The acoustic records of anchovy abundance were
compared with the distribution of the sea surface
temperature, as well as with the strati®cation of the
water column. To quantify that strati®cation, the /
 196 J.E. Hansen et al.

parameter of stability (Simpson, 1981) was used Figure 2. Annual values of total anchovy abundance (sa) at
(in J m±3) as a measure of the work needed to mix the the grid nodes (empty circles), and mean of these values
water column. The parameter is de®ned as follows. (dark circles) as a function of the respective acoustic biomass
estimates.
Z0
/ ˆ gh …q ÿ q0 †z dz
ÿh

where:
g is gravity acceleration,
h is total depth,
q is density,
q0 is mean density of the column and
z is depth.
Two vertical sections per survey were selected, one
off the ValdeÂs Peninsula (42°30¢S) and the one other
(43°20¢S) about the latitude of Escondida Island, to
examine the relative abundance of anchovy shoals by
water layers according to the thermal structure Thereafter, anchovy abundance at the grid nodes
(Transects A and B, Fig. 1). was computed as a function of their corresponding
values: either SST (class interval, 0.5°C) or stability
Interpolation of the original data onto grids index. The latter were split into four categories, or
The raw data on anchovy abundance (sa), sea surface stability classes, as follows.
temperature (SST) and stability index (/) from each
(i) Stability values £ 9.99, representing zones where
year were used as input for the software Surfer 6.01
the water column was homogeneous.
(Golden Software, 1995) to produce regularly spaced
(ii) Values between 10 and 69.99, which were asso-
arrays of values corresponding to the same geograph-
ciated with the presence of thermal fronts, as the
ical locations as the original from the irregularly
mean value of this range, i.e. the stability line of
11 spaced latitude±longitude recordings, by kriging. The
40 J m±3 was used by Martos and SaÂnchez (1997)
same grid geometry (24 ´ 29 km) was produced as
to de®ne the mean position of the fronts.
output for every year and for all three kinds of data.
(iii) Values between 70 and 129.99, corresponding to
This procedure was analogous to that used by Swain
de®nitively stable zones, which are located rel-
(1999) to compare the distribution of bottom tem-
atively close to the fronts.
peratures and cod abundance in the southern Gulf of
(iv) Stability index ³ 130, representing highly stable
Lawrence. In the present study, the goodness of ®t of
marine zones, which are located far away from
the interpolated, grid-node values to the original ones
thermal front axis.
was evaluated by analysing the residuals for every
annual grid (sa, SST, stability), and neither trends nor To look for any relationship between the environ-
signi®cant distortions were found. mental variables and the anchovy abundance data
Figure 2 shows also that both the total abundance from the four surveys together, the values of ®sh
by year (sums of anchovy sa at the grid nodes) and density at the grid nodes (sai) were standardized by
their means were fair indicators of the respective sur- taking into account their annual mean (…sa ; y†) and
vey biomass as estimated from the original acoustic the greatest annual mean, which was that of the year
values. The lack of independence between the two sets 1995 …sa ; 1995†, as follows:
of abundance values is evidence of the agreement
resulting from the kriging method, even though the sa i; y ˆ …sa i, y†  …sa ; 1995†=…sa ; y†
anchovy distribution is patchy. Nevertheless, the high for y ˆ 1993; . . . ; 1996
coef®cients of determination included in the ®gure
may be not very meaningful, because they are largely These standardized values were also tested for
due to the occurrence of two clusters of values for each relations to the above-mentioned categories of both
data type. SST and stability.

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 Distribution of Argentine anchovy and sea temperature 197

RESULTS shows the average of the annual percentages of ®sh

Distribution and abundance of anchovy during the surveys
Anchovy were found on most of the acoustic transects
during the 4 years of the survey, mainly on the inner
and intermediate shelf up to 70 m deep. Even though
the estimation of the absolute abundance of the spe-
cies is not the main aim of this paper, Table 1 indi-
cates the estimated biomass by survey, and the
coef®cients of variation. The magnitude of the inter-
annual oscillations of abundance agree with those
observed both in this and other anchovy stocks
(Ciechomski and SaÂnchez, 1988). Size sampling
showed that ®sh ranged from 78 to 199 mm in total
length, with little variation between years. Figure 3
according to size within the ®ve strata. Most of the
anchovies at almost every size were found in Stratum 2,
in front of the ValdeÂs Peninsula, i.e. between 42 and
43°S. One constant pattern was the occurrence of the
smallest ®sh at the mouth of San MatõÂas Gulf
(Stratum 1), but they were not exclusive to that zone.
They were immature, as the size at 50% maturity has
been estimated to be » 120 mm (Hansen and Madir-
olas, 1996). On the other hand, the highest abun-
dances of medium-sized and larger ®sh (> 125 mm)
were found in different regions from one year to
another. Within the Strata 3 and 4, larger anchovies
prevailed, whereas medium sized ®sh were often found
in the southernmost stratum.

Distribution of sea temperatures during the surveys

Table 1. Estimated values of anchovy biomass and coef®-
Three frontal zones have been identi®ed: (i) at the
cient of variation, by survey.
mouth of San MatõÂas Gulf; (ii) in front of the ValdeÂs
Year Estimated biomass (t) Coef®cient of variation Peninsula; and (iii) along the Patagonian coast, close
to Escondida Island (Fig. 4). Within the area corres-
1993 764 710 13% ponding to the thermohaline front off San MatõÂas
1994 975 298 19%
Gulf, the maximum surface temperatures (» 16°C)
1995 1969 881 11%
were recorded in the northern part, whereas the
1996 2198 822 20%
minimum values (» 13°C) corresponded to the zone

Figure 3. Average of the annual percentages of ®sh according to size within the ®ve strata.

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 198 J.E. Hansen et al.

Figure 4. Sea surface temperatures and relative abundance of Patagonian anchovy (from acoustic measurements, m2/nm2), as
observed during 1993±1996.

closer to the ValdeÂs Peninsula. The horizontal
gradients in this front were greater during 1995 and
1996 (0.030 and 0.025°C km±1) than in 1993 and
1994 (0.015 and 0.020°C km±1).
Within the frontal zone off the ValdeÂs Peninsula,
minimal surface temperatures (12±13°C) were recor-
ded from the coastal side of the front, whereas the
highest values (13.5±16°C) corresponded to the sur-
face layer of the strati®ed side. The sharpest gradients
of SST (0.020 and 0.025°C km±1) were found in 1995
and 1996. Finally, the front along the Patagonian
12 coast south of the ValdeÂs Peninsula, even though it
was closer to the coast than the others, showed similar
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features, i.e. minimal temperatures on the coastal side
(12±13°C) and maximum values towards the strati®ed
region (13.5±15°C). The front had a weak manifes-
tation in the surface during 1995, and reached its
maximum gradient in 1996 (0.035°C km±1).
There were some interannual changes in the mean
position of the fronts (Fig. 5). In 1993 and 1994, the
axis of the front off the ValdeÂs Peninsula and that of
13 the one south of 43°S were located » 100 km from the
coast. Both fronts were closer to the continent during
the following years, particularly that off Escondida
Island (20 km in 1995, and 28 km in 1996), when the
highest values of stability index, and the widest

Figure 5. Values of the stability index and relative abundance of Patagonian anchovy, as observed during the different surveys.
water-strati®ed areas were found. The position of the
front off San MatõÂas Gulf was similar during the
4 years, showing a maximum variation of » 10 km.
The distributions of temperature (Fig. 4) and stability
(Fig. 5) in San MatõÂas Gulf look orthogonal at times if
they are superposed, e.g. in 1994, because this is a
thermohaline front not a tidal one.
Vertical sections at the latitudes of the ValdeÂs
Peninsula (Fig. 6) and Escondida Island (Fig. 7) have
shown that those tidal fronts had also bottom mani-
festations (particularly strong off Escondida Island
during 1995), which were farther from the coast than
the manifestations at the surface. The ®gures show
typical sections, where frontal regions separate coas-
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Distribution of Argentine anchovy and sea temperature 199
tal, homogeneous zones from strati®ed, outer regions.
The strati®ed zone of both fronts had a similar
structure, consisting of two homogeneous layers with
a thermocline layer in between, from 20 to 40 m
deep. The temperatures within the coastal region
ranged from 12 to 13°C over the 4 years, whereas
within the strati®ed part they oscillated between 13.5
and 16°C at the upper layers and 8±10°C in the
lower layers.
Sea surface temperature and presence of anchovy
The species occurrence, as acoustically determined by
integrating echo values from the total water column,
was ®rst graphically associated with sea surface
200 J.E. Hansen et al.
isotherms (Fig. 4). Notice that the biggest squares,
representing the highest records of anchovy abun-
dance, were related to the presence of the sharpest
gradients of sea surface temperature, » 0.02°C or
more per km, as indicated by the relative proximity of
the isotherms.
Within the area corresponding to the thermohaline
front off San MatõÂas Gulf, the highest anchovy densities
were recorded in 1995 (total sa ˆ 371 181 m2/nm2) and
1996 (221 379), i.e. when high horizontal gradients
were found (Martos and SaÂnchez, 1997). The results of
the four surveys support previous observations from the
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Figure 6. Vertical pro®les off the ValdeÂs
Peninsula (42°30¢S) showing the distri-
bution of Patagonian anchovy density
values (squares, see the sa scale at the
bottom), as observed during the surveys
carried out in 1993 (A), 1995 (B), and
1996 (C). Arrows and station numbers
show the sailing course. Suns and moons
23 denote daytime and night, respectively.
1993 cruise (Hansen and Madirolas, 1996), as the
thermal gradient of SST off San MatõÂas Gulf has
shown to be a favourable zone for small anchovies.
Great concentrations of anchovy were always found
within the frontal zone off the ValdeÂs Peninsula. They
were maximal in 1995 (total sa ˆ 319 856 m2/nm2)
and 1996 (407 230), when the sharpest gradients of
SST were veri®ed (Martos and SaÂnchez, 1997).
High ®sh densities, which were maximal during 1995
(total sa ˆ 421 163 m2/nm2) and 1996 (393 795), were
also observed along the Patagonian coast south of the
ValdeÂs Peninsula. In 1995, this sea front had a weak
 Distribution of Argentine anchovy and sea temperature 201

Figure 7. Vertical pro®les near Escondida Island (» 43°20¢S) showing the distribution of Patagonian anchovy density values
(squares, sa scale at the bottom), as observed during the surveys carried out in 1995 (A) and 1996 (B). Station numbers show the
sailing course. Suns and moons denote daytime and night, respectively.

manifestation at the surface, but the sharpest gradients
of water temperature occurred at the sea bottom. The
surface manifestation of the front reached its maximum
in 1996 (Martos and SaÂnchez, 1997).
The results of pooling by survey the anchovy
abundance at grid nodes as a function of the SST class
at the same positions are represented in the Fig. 8A±C.
The thermal bounds were 11 and 17°C, with a pre-
ferential range between 12.5 and 16°C, even though a
pair of very high values was located at SST of 16.5 and
17°C during the 1996 survey (Fig. 8C).
Within the above-mentioned limits, the absolute
value of SST did not seem to determine the massive
occurrence of Patagonian anchovy. In different years,
the frequency of observations (Fig. 8A) was distri-
buted around different mean SST values, i.e. 14°C in
1993 and 1994, 13.5°C in 1995, and 15°C in 1996.
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The total abundance by SST class re¯ected the same
trend (Fig. 8B) because none of the well-represented
classes had a very high mean abundance.
The same conclusion arises from pooling all the
standardized abundance values (Fig. 9A±C). The fre-
quency distribution (Fig. 9A) is approximately nor-
mal, around the range of 13.5±14°C, and none of the
SST classes showed an especially high mean, except
those of 16.5 and 17°C, which corresponded only to a
single pair of values, as mentioned before.
Anchovy distribution and location of the sea fronts
A ®rst view of the horizontal distribution of anchovy
abundance values for the whole water column showed
that the species was found both in thermally strati®ed
and mixed areas (Fig. 5). Nevertheless, the shoals
seemed to be mainly found near the mean location of
202 J.E. Hansen et al.
Figure 8. Annual anchovy abundance (sa) by sea surface
temperature intervals. A, frequency distributions, i.e. fraction
of the total area associated with each temperature; B, total
abundances (m2/nm2); C, mean abundances (m2/nm2).
the fronts, which appeared to be the axis for the dis-
tribution. The ®gure shows that the above-mentioned
annual changes in the position of the tidal fronts
agreed with some variations observed in the location
of the main concentrations of adult anchovy, which
were found closer to the continent during 1995 and
1996 than they were during the previous surveys.
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Figure 9. Standardized anchovy abundance (sa*i, y) by
sea surface temperature intervals. A, frequency distributions;
B, total abundances (m2/nm2); C, mean abundances
(m2/nm2).
The results of comparing the anchovy abundances
at the nodes with the stability of the water column
at the same positions are shown in Fig. 10A±C.
Figure 10A shows the number of sa,y-values
(y ˆ 1993,¼,1996) within each stability class. As the
grid is regular, that number is proportional to the total
marine area of each stability category during the
respective cruise. It is noteworthy that the area of
homogeneous conditions of the water column (Class 1
stability) was rather constant throughout the whole

Figure 10. Annual anchovy abundance (sa) by stability class.
A, frequency distributions; B, total abundances (m2/nm2); C,
mean abundances (m2/nm2).
period, and a bit less than that in the areas corres-
ponding to the other three categories. In the 1993 and
1994 surveys, the most represented categories were 2
(zones of thermal front in¯uence) and 3. On the other
hand, most of the data from the 1995 and 1996 cruises
corresponded to the category 4, which showed a wide
extension.
The zone of the fronts was generally the most
important with respect to the total abundance at the
grid nodes (Fig. 10B), even though during the last two
surveys the total occurrence of anchovy within both
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Distribution of Argentine anchovy and sea temperature 203
the homogeneous zone (1) and the most stable one (4)
was also signi®cant.
The most relevant of our results, in our view, is that
the mean abundance by stability class indicated, for
every survey, that the zones in¯uenced by thermal
fronts contained the most dense anchovy concentra-
tions (Fig. 10C). Most of the time (the 1994 cruise
being the exception) those zones did not represent the
widest of the surveyed areas, as mentioned before. The
means within the homogeneous, coastal zones were
rather high and followed the respective trend of the
total values. On the other hand, the means corres-
ponding to stable areas were de®nitively lower, even in
the surveys when the total abundance within these
zones was important (years 1995 and 1996). These
results indicate the occurrence of many low-density
®sh shoals.
The standardized abundance data (Fig. 11A±C)
suggest that the four stability classes are well repre-
sented. As mentioned before, the mean abundance
within the homogeneous class is high. But both the
mean and the total abundance had the greatest values
within the areas associated with thermal fronts.
Anchovy distribution and vertical thermal structure
The vertical pro®les at the latitudes of the ValdeÂs
Peninsula (Fig. 6A±C) and Escondida Island (Fig. 7A
and B) show anchovy shoals within the range
8.5±16.5°C. Nevertheless, most of those inhabiting
> 14°C were recorded at night, when ®sh usually
occupy a water layer near the sea surface, even though
they could not be detected within the upper 5.5 m.
The highest ®sh concentrations in the daytime were
found in waters ranging from 10.5 to 13.5°C, whereas
they ranged from 13 to 16°C at night.
During the day, anchovies seemed to prefer the
thermocline layer or the one immediately above it
(Fig. 6B and C), whereas only a few ®sh shoals were
located below the thermocline (Fig. 6A).
Another kind of distribution observed in these
surveys consists of some ®sh shoals inhabiting a
coastal region of vertically homogeneous waters
where a thermal horizontal gradient is present at the
surface (Fig. 7A). Under these conditions, the
15 biomass in coastal regions south of 43°S can be high
(Fig. 13A). It can also be high (as seen during the
1996 survey) when the sea front is very close to the
coast and no homogenous zone is present at that
latitude (Fig. 7B). As a result, abundant anchovy
were often found near either surface or bottom
manifestations of the front.
The above-mentioned patterns of Patagonian
anchovy distribution in relation to sea temperatures
204 J.E. Hansen et al.
Figure 11. Standardized anchovy abundance (sa*i, y) by
stability class. A, frequency distributions; B, total abun-
dances (m2/nm2); C, mean abundances (m2/nm2).
16 eous coastal waters south of 43°S when a thermal
and stability can be regarded as valid for the main
concentrations of the species during all the surveys.
Though more complicated images were observed on
some occasions (Fig. 7A), no relationship existed
between thermal structure and the mean size of ®sh in
trawls. In that sense, the only clear trend observed was
the presence of small adult anchovies off San MatõÂas
Gulf, where (as mentioned before) the highest sea
surface temperatures were recorded. Castello (1997)
has also pointed out that small anchovies from
southern Brazil preferred the warmest waters.
Ó 2001 Blackwell Science Ltd., Fish. Oceanogr., 10:2, 193±206.
DISCUSSION
In conclusion, Patagonian anchovies were present on
almost every acoustic transect during the four surveys,
mainly on the inner and intermediate shelf 70 m deep,
and remarkably in front of the ValdeÂs Peninsula
(42±43°S). The occurrence of the smallest ®sh at the
mouth of San MatõÂas Gulf was also a constant feature,
whereas the location of medium-sized and larger ®sh
differed from year to year.
The lower and upper sea surface temperature
bounds for anchovy distribution were 11 and 17°C,
with a preferential range between 12.5 and 16°C, but
no absolute value of SST could be indicated as the
most favourable for the species. Our results con®rm
and complement the preliminary analysis concerning
the main oceanographic features of the area performed
by Martos and SaÂnchez (1997). Although anchovies
were found either in thermally strati®ed or mixed
areas, their annual distributions were clearly related to
the three frontal zones occurring in the region during
late spring to early summer, and they changed
accordingly to the mean position of the fronts in
different years. Both total abundance and mean
abundance within zones of thermal front in¯uence
were usually the most important, and most of the high
anchovy densities were usually recorded within the
areas showing the sharpest gradients of sea surface
temperature. The bottom manifestations of the fronts
were also sometimes important in that sense.
Vertically, anchovy were distributed within the
temperature range of 8.5±16.5°C, with an apparent
preferential range between 10.5 and 13.5°C during the
daytime. During several research surveys carried out
on the southern Brazilian shelf, Castello (1997) has
found adult anchovies inhabiting waters between
9.5 and 21.7°C. In Patagonian waters, ®sh seemed to
prefer the layer above the 20±40 m-deep thermocline,
but they were also abundant in vertically homogen-
horizontal gradient was present at the surface.
In close agreement with the results presented in
this paper, some other studies of seasonality and
geography of anchovy and sardine spawning activity
have shown that temperature itself, provided it falls
within the species tolerance, may be less important in
de®ning favourable seasons and/or locations for
spawning than the occurrence of mechanisms pro-
moting retention of larvae near the coast while min-
imizing turbulent mixing of the water column (Bakun
and Parrish, 1991). Accordingly, Green Ruiz and
Hinojosa Corona (1997) have pointed out that the
patterns of water masses and temperature fronts in the

Gulf of California, as derived from satellite images,
could be related to the distribution and abundance of
northern anchovy (Engraulis mordax) eggs. In a per-
haps wider context, Castillo et al. (1996) have con-
cluded that the distributions of three pelagic resources
from northern Chile (anchovy, Engraulis ringens; sar-
dine, Sardinops sagax; and jack mackerel, Trachurus
murphyi) were associated with the occurrence and
intensity of thermal fronts. In particular, the Southern
Paci®c anchovy was found to be associated with the
anterior zone of a thermal front, and pushed towards
the coast during the seasons when the front became
stronger.
Bakun and Parrish (1991) have described the
mechanisms that relate to the Patagonian sea fronts
and lead to positive conditions for anchovy repro-
duction in late spring. They include strong tidal
mixing over the shallower areas, carrying nutrients
upwards to support both photosynthetic activity
(maximum at this time) and production of food par-
ticles which then ¯ow into a zone of stability, i.e.
the mid-depth thermocline layer.
By using the parameter of stability to de®ne the
location of the frontal regions objectively during
the spawning season of the stock, our results con®rm
the af®nity of mature or potentially mature anchovies
for those areas. The implications for improving the
allocation of commercial ®shing effort and manage-
ment are obvious. It is noteworthy that small, still
immature ®sh like those found off San MatõÂas Gulf
were also more abundant in frontal zones. The results
might suggest that the preference of the species for
adequate spawning habitats is established prior to
sexual maturity itself.
Even though the sea fronts are the main zones for
Patagonian anchovy spawning, they are not the
unique reproductive grounds. The spawning areas were
estimated to range over as much as 54±64% of the
whole surveyed area in the cruises carried out between
1993 and 1995 (SaÂnchez et al., 1996). On the same
cruises, egg data and adult data did not show exactly
the same pattern. Accordingly, our observations show
that high concentrations of ®sh can also be found
either in mixed or strongly strati®ed waters during this
season. It is also known that adult, spent or resting
anchovies are present in the region during other sea-
sons, after the tidal fronts have been eroded. In late
summer and early autumn, when ®sh feed intensely on
the increased zooplanktonic biomass resulting from
the previous phytoplanktonic bloom, anchovy distri-
bution on the Patagonian shelf, although still patchy,
was found to be more dispersed (Angelescu and
Anganuzzi, 1981).
Ó 2001 Blackwell Science Ltd., Fish. Oceanogr., 10:2, 193±206.
Distribution of Argentine anchovy and sea temperature 205
ACKNOWLEDGEMENTS
This paper is INIDEP contribution number 1156.
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