INTRODUCTION
Studies dealing with the age determination and
growth of fish species are considered a crucial
topic in fisheries research (Hoenig and Gruber
1990; Officer et al. 1996; Campana 2001), since
they provide basic information for stock assess-
ment (Gallagher and Nolan 1999).
Brazilian flathead Percophis brasiliensis Quoy
and Gaimard 1825 (Suborden Trachinoidei) is a
MARINE AND FISHERY SCIENCES 32 (2): 103-114 (2019). https://doi.org/10.47193/mafis.3222019121902
FIRST DATA ON THE AGE AND GROWTH OF BRAZILIAN FLATHEAD
Percophis brasiliensis (PISCES: PERCIFORMES)
IN SAN MATÍAS GULF, NORTHERN PATAGONIA (ARGENTINA)
MARÍA E. LÓPEZ1, 2, 3, MATÍAS OCAMPO REINALDO2, 4, 5, KARINA A. RODRÍGUES2, 6,
M. RITA RICO6, MARGARITA B. SÁEZ6and RAÚL A. GONZÁLEZ2, 4, 5
1Instituto de Desarrollo Costero “Dr. H. C. Héctor E. Zaixso” (IDC), Universidad Nacional de la
Patagonia San Juan Bosco (UNPSJB), Ruta 1 s/n Km 4, 9000 Comodoro Rivadavia, Argentina
e-mail: melopez@live.com.ar
2Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina
3Centro de Investigación y Transferencia “Golfo San Jorge”
(CIT-GSJ), CONICET-UNPSJB-UNPA, Ruta 1 s/n Km 4, 9000 Comodoro Rivadavia, Argentina
4Centro de Investigación Aplicada y Transferencia Tecnológica en Recursos Marinos “Almirante
Storni” (CIMAS), Güemes 1030, 8520 San Antonio Oeste, Argentina
5Escuela Superior de Ciencias Marinas (ESCiMar), Universidad Nacional del Comahue (UNCo),
San Martín 224, 8520 San Antonio Oeste, Río Negro, Argentina
6Instituto Nacional de Investigación y Desarrollo Pesquero (INIDEP),
Paseo Victoria Ocampo N° 1, B7602HSA - Mar del Plata, Argentina
ABSTRACT. Brazilian flathead Percophis brasiliensis is a commercially important perciform fish caught as bycatch
by trawlers throughout its latitudinal distribution range (from 23° S in Brazil to 47° S in Argentina). It is associated
with the catches of Argentine hake Merluccius hubbsi and Patagonian red shrimp Pleocticus muelleri in San Matías
Gulf (SMG). The main objective of this study was to describe the first data on the growth of this species in SMG and
to compare the results with available information for other areas of the Argentine Continental Shelf (ACS). The study
was carried out using 294 specimens (196 females, 97 males and one of indeterminate sex) collected over four non-
consecutive years from commercial/recreational fishing and during research cruises. The age of individuals was deter-
mined by reading and counting opaque-hyaline rings on the sagittae otoliths. The maximum ages observed were 14
years for females and 12 years for males. Growth parameters were estimated using the von Bertalanffy mode.
Significant differences in L∞parameter between sexes were found (L∞males =54.58 cm; L∞females =78.31 cm,
p< 0.05). Growth parameters were different from those determined in previous studies for the ACS.
Key words: Age determination, growth curve, Percophis brasiliensis, gulf region, poor-data species.
103
coastal fish species of demersal-benthic habits.
Its distribution extends from 23° S (Rio de
Janeiro, Brazil) to 47° S (north of Santa Cruz
Province, Argentina), and from the coast down to
depths of 75 m (Cousseau and Perrotta 2013). P.
brasiliensis is mainly caught by bottom trawl nets
in the context of a multi-species fishery in the
coastal ecosystem of Buenos Aires Province
(CEB) on the Argentine Continental Shelf (ACS)
(Carozza et al. 2001). Several studies on the com-
position of catches and growth were carried out in
this area (Carozza et al. 2018; Rico et al. 2018).
This species mainly inhabits the northeast area of
San Matías Gulf (SMG) between the coastline
and the 90 m isobath (López et al. 2012). It is
caught as bycatch in a multispecific fishery tar-
geting Argentine hake Merluccius hubbsi
(Romero et al. 2013) and in a recent fishery tar-
geting Patagonian red shrimp Pleoticus muelleri
(Sepúlveda 2018). Landings recorded since the
early 90s have always amounted to a few tons
(8.75 ± 21.92 SD), except for a peak recorded for
the period 1999-2000 (82.3 ± 33.66 SD).
At present, Brazilian flathead is considered a
data-poor species in SMG. Biological information
on P. brasiliensis in SMG fishery is scarce and
insufficient for inferring the status of its population
structure and dynamics. However, some biological
aspects of this species were previously studied in
the CEB, where the peak of reproductive activity
was observed in November and gonads in resting
stage were detected in May, June, and August
(Rodrigues et al. 2010, 2013). P. brasiliensis is a
long-living, slow-growing species with maximum
ages ranging from 15 years for females to 19 years
for males (Rico and Sáez 2010; Barretto et al.
2011; Sáez et al. 2011). Differential growth
between the sexes from the first year of life has
been observed with females reaching greater sizes
than males of the same age (Barretto et al. 2011).
Despite their longevity, females mature in the sec-
ond year (35.6 cm total length, TL) and males in
the first (25.5 cm TL) (Rodrigues 2012). The
chemical composition of otoliths also highlighted
a possible difference between individuals belong-
ing to SMG and the Argentine-Uruguayan Com-
mon Fishing Zone (AUCFZ) (Braicovich and Timi
2008; Avigliano et al. 2015). This evidence may
indicate that P. brasiliensis in SMG could be a rel-
atively isolated subpopulation or stock, as
observed for other species of similar latitudinal
distribution range, such as Argentine hake M.
hubbsi (Sardella and Timi 2004; Ocampo Reinaldo
et al. 2013) and the Brazilian sandperch Pinguipes
brasilianus (Timi et al. 2008).
The aim of this study was to determine first
data on the age and growth parameters of P.
brasiliensis in SMG as an initial step towards
understanding basic aspects of the biology and
population dynamics of this species and its role in
demersal and coastal fish assemblages.
MATERIALS AND METHODS
Samples were collected over four non-consecu-
tive years from three sources: research cruises,
commercial landings and coastal rod fishing
(Table 1). Total length (TL) to the nearest cen-
104 MARINE AND FISHERY SCIENCES 32 (2): 103-114 (2019)
Table 1. Source and number of samples. RC: research cruis-
es, CL: commercial landings from the Patagonian red
shrimp and the Argentine hake fisheries, RF: rod
fishing.
Month 2009 2011 2013 2014
February - - - 3 (CL)
March - - - 18 (CL)
April - - - 5 (CL)
May 25 (CL) - - -
June 17 (CL) - 3 (CL) -
August 14 (CL) - - -
November 200 (RC) 9 (RF) - -
Total 256 9 3 26
timeter, total weight (TW) in grams and sex were
recorded for each fish. Sagittae otoliths were
extracted and one otolith from each pair was
washed, dried and embedded in opaque epoxy
resin. The otoliths were sectioned transversally
through the nucleus region in order to obtain a 0.5
mm-thick section using a Maruto MC-201 micro-
cutter.
Age validation
In order to validate the periodicity of the
growth rings, the type of band (opaque-translu-
cent) on the edges in the otolith sections was
recorded.
Age determination
To determine the age of the individuals, the
pairs of opaque-translucent bands in the otolith
sections were counted by two independent readers
under incident and transmitted light using a stere-
omicroscope at 40X magnification (Figure 1). Age
was determined to the nearest lower according to
Barretto et al. (2011) without knowledge of the
length and sex of the specimen, considering that
one opaque and one translucent band representing
one year of the fish life. The average percent error
(APE) (Beamish and Fournier 1981) and the coef-
ficient of variation (CV) (Chang 1982) were used
to validate the age determination of both observers.
Growth
The von Bertalanffy (1938) growth model
(VBGM) was used to characterize the growth in
length as a function of age using the following
equation:
where TLtis the average total length (cm) at age t
eTL t
=LE(1 )-+
¥0
t
-
(- ))kt(
t
105
LÓPEZ ET AL.: GROWTH OF P. BRASILIENSIS IN SAN MATÍAS GULF
Figure 1. Thin section of the saggitae otolith of a 4-years-old female Percophis brasiliensis. N: nucleus, O: opaque band, H:
hyaline band; the numbers refers to the annual rings.
(years), L∞is the maximum total length (cm), kis
a growth rate parameter (year-1), t0is the theoret-
ical age (years) at zero length and Etwith Normal
distribution since we were working with averages
TLt.
The model was fitted using the generalized
least squares method (Kimura 1980). Growth
parameters (L∞, kand t0) were estimated using the
maximum likelihood method (Cerrato 1990).
Comparisons between growth curves of both
sexes were carried out using the likelihood ratio
and the χ2 distribution ( p <0.05; Cerrato 1990).
RESULTS
Age determination
Age was determined for a total of 294 individ-
uals (196 females, 97 males and one undefined).
The average percent error (APE =0.28%) and the
coefficient of variation (CV =0.40%) indicated
that the criteria for age determination was consis-
tent across the readers (Cerrato 2000).
TL values ranged from 33 to 71 cm for females
and 32 to 63 cm for males. Both sexes showed
unimodal frequency distribution at 55 cm for
females and 50 cm for males (Figure 2).
Age validation
Both bands (opaque and hyaline) were present
in most of the months analyzed (Figure 3). The
opaque band was frequent in February, although
deposition could have started months earlier. In
addition, the hyaline ring was deposited from
March to August.
Growth
Ages ranged from 2 to 14 years for females and
3 to 12 years for males, with a mode of 4-year-old
specimens in both sexes (Figure 4; Table 2). The
VBGM showed a good fit to the observed data
with females larger than males at the same age
(Figure 5). When comparing the three parameters
together (L∞, k, t0), significant differences were
found ( p <0.05) between the growth curves by
sex. Comparisons of the parameters individually
detected significant differences for the asymptotic
length only (Table 3).
106 MARINE AND FISHERY SCIENCES 32 (2): 103-114 (2019)
Figure 2. Frequency distribution of total length by sex of Percophis brasiliensis.
0
5
10
15
20
25
30
35
40
32 36 40 44 48 52 56 60 64 68 72
Frecuency
Total length (cm)
Males
Females
DISCUSSION
The accuracy of age readings and the low per-
centage of otoliths discarded due to malforma-
tions show that the sagittae otoliths of P.
brasiliensis appear to be suitable for reading ages,
like most sagittae otoliths of temperate-water fish
(Panella 1974; Pauly 1980; Lombarte and
Lleonart 1993; Morales-Nin 2000). In all the thin
sections analyzed the nature of the first ring was
opaque, coinciding with observations for the
same species on Buenos Aires coast (Barretto et
al. 2011). This could be because the opaque band
is formed in the warm months (late spring and
summer) when recruitment of individuals to the
populations of P. brasiliensis on the Argentine
coast is recorded (Rodrigues 2012). On the other
hand, the hyaline ring corresponds to the coldest
seasons (autumn and winter). Although the vali-
dation was carried out by mixing months of dif-
107
LÓPEZ ET AL.: GROWTH OF P. BRASILIENSIS IN SAN MATÍAS GULF
Figure 3. Monthly percentage of opaque and hyaline edges in whole otoliths of Percophis brasiliensis in SMG.
Figure 4. Frequency distribution of age by sex of Percophis brasiliensis.
0
20
40
60
80
100
JASONDJFMAMJ
Edge proportion (%)
Month
Opaque Hyaline
n = 3
n = 14 n = 212 n = 1 n = 5 n = 2 n = 2
0
5
10
15
20
25
234567891011121314
Frequency (%)
Age (years)
Males
Females
108 MARINE AND FISHERY SCIENCES 32 (2): 103-114 (2019)
Table 2. Range (cm), mean total length (TL, cm) per age (in years) of females, males and pooled data of Percophis brasiliensis
from SMG. SD: standard deviation.
General Females Males
Age Range Mean TL ± SD (n) Range Mean TL ± SD (n) Range Mean TL ± SD (n)
2 35-43 37.6 ± 2.97 (8) 35-43 37.6 ± 2.97 (8) -
3 32-48 40.0 ± 3.85 (26) 35-48 40.9 ± 3.55 (22) 32-46 39.00 ± 4.11 (16)
4 33-53 44.7 ± 3.84 (66) 33-53 45.2 ± 3.92 (45) 34-49 43.50± 3.47 (21)
5 35-56 48.1 ± 4.26 (49) 39-56 49.4 ± 3.78 (33) 35-54 45.60± 4.15 (16)
6 42-61 51.3 ± 4.24 (40) 45-61 53.5 ± 3.44 (24) 42-53 48.20± 3.25 (16)
7 49-63 54.7 ± 3.77 (19) 51-60 55.8 ± 2.59 (12) 49-63 52.70± 4.82 (7)
8 47-65 55.8 ± 4.71 (23) 52-65 58.4 ± 3.58 (15) 47-53 51.00 ± 1.85 (8)
9 52-70 59.2 ± 4.60 (26) 54-70 60.6 ± 4.08 (20) 52-54 53.40± 0.89 (5)
10 52-67 58.6 ± 4.94 (13) 55-67 60.3 ± 4.92 (9) 52-57 54.75 ± 2.06 (4)
11 51-67 60.1 ± 5.25 (8) 59-67 63.2 ± 3.27 (5) 51-57 55.00 ± 3.46 (3)
12 51-61 56.0 ± 7.07 (2) 61 (1) 51 (1)
13 71 (1) 71 (1) -
14 68 (1) 68 (1) -
Figure 5. Growth curves of Percophis brasiliensis by sex from San Matías Gulf. Bars represent standard deviation. VBGM: von
Bertalanffy growth model.
0
10
20
30
40
50
60
70
80
0246810121416
Total length (cm)
Age (years)
VBGM females VBGM males
Average of females observed Average of males observed
ferent years without having the entire cycle and
with few individuals in some months, the periods
of growth ring formation coincided with those
reported by Barretto et al. (2011) for individuals
from Buenos Aires coast. Similarly, the formation
of rings at similar times for fish that inhabit
Southwestern Atlantic coast has been described.
Scartascini et al. (2015) studying the seasonality
of the otoliths of the white croaker Micropogo-
nias furnieri in SMG reported the deposition of
opaque bands between November and January,
coinciding with that reported for weakfish
Cynoscion guatucupa (López Cazorla 2000) in
Bahía Blanca waters and for wreckfish Plyprion
americanus in southern Brazil (Peres and
Haimovici 2004).
Considering that the first pair of rings are usu-
ally harder to identify than the subsequent pairs,
some authors have suggested not counting it
(Peres and Haimovici 2004). In order to compare
the estimates made, this study used the criteria
described by Barretto et al. (2011) to estimate the
age of the species in another site of its distribution.
The maximum ages observed in this study (14
years for females and 12 for males) were higher
than the maximum ages previously recorded in
CEB for this species: 6 years (Tomo 1969), 7
years (San Román 1974) and 12 years (Perrotta
and Fernández-Giménez 1996). However, they
were lower than those recorded by Barretto et al.
(2011): 15 years for females and 19 years for
males (both between 60 and 70 cm TL). Individ-
uals belonging to the 0 and 1 year age classes
were not represented in this study even though the
trawl nets used during the research cruises had a
small mesh (40 mm) inner cover. Other studies
carried out in CEB also reported the absence or
low occurrence of individuals belonging to these
year classes (Rico et al. 2018). Considering that
small individuals inhabit mainly shallower areas
(Barretto 2007) their presence would not have
been detected in this study because these areas
were not sufficiently surveyed.
The VBGM was adequate for describing the
growth of P. brasiliensis. Differential growth
between sexes was observed in this study with
females larger than males at the same age as
described previously by San Román (1974) and
Barretto et al. (2011). P. brasiliensis is a slow
growing, relatively long-lived species like other
perciform fishes studied in SMG (Rubinich and
González 2001; González 2006).
The estimated L∞for females (78.31 cm TL)
was higher than that previously reported (65.2 and
63.5 cm TL) in the northern area of distribution
(CEB) (Barretto et al. 2011), while L∞for males
(54.58 cm TL) was lower than the estimated for
the species outside the SMG (58.1 cm and 58.7
cm TL, Barretto et al. 2011). Although habitat dif-
ferences between the two areas (Warm Tempera-
ture Southwestern Atlantic province versus the
“Magellanic Province”) (Spalding et al. 2007)
might affect the growth of these species, differ-
ences in L∞could also be partially linked to the
quality of the data since Barretto et al. (2011) used
a wider range of age classes in their study (17 age
classes versus 14 in this study). In our study, the
lack of individuals smaller than 33 cm TL might
explain the relatively high values of t0and conse-
quently have an effect over the grow rate parame-
ter (k). However, differences in TL between males
and females within SMG population compared to
other stocks were clear. Differences could also be
explained by the different fishing intensity to
109
LÓPEZ ET AL.: GROWTH OF P. BRASILIENSIS IN SAN MATÍAS GULF
Table 3. von Bertalanffy’s growth parameters (L∞,k,t0) for
Percophis brasiliensis from San Matías Gulf. The
general curve includes one individual of indetermi-
nate sex shown as comparison purpouses with other
studies.
L∞(cm) k (year-1) t0 (years) N
General 86.06 0.08 -5.51 294
Females 78.31* 0.11 -3.61 196
Males 54.58* 0.36 -0.46 97
*Indicates significant differences (p< 0.05).
which the population of Brazilian flathead has
been subjected at both sites (CEB and SMG). The
CEB fishery began around 1960 and catches
increased up to 8,350 t in 1997. Although catches
have decreased since then, more than 7,000 t of
Brazilian flathead are still landed in CEB (Rico et
al. 2018). The intensity and long-term fishing
pressure of the CEB fishery could have caused the
removal of adult individuals resulting in a greater
relative abundance of individuals between 3 and 5
years old (Rico et al. 2018).
The estimated k (0.11 year-1) and t0(-3.61
years) for females were lower than those calculat-
ed previously for Buenos Aires stock by Barretto
et al. (2011) (k =0.29 year-1 in spring and 0.26
year-1 in winter, t0=-1.15 years in spring and
-2.01 years in winter). However, if data from
smaller (<33 cm TL) and larger individuals (>70
cm TL) were added to the VBGM the estimated
value of t0could be lower, leading to a higher
value of k. On the other hand, parameters of the
VBGM in males showed more realistic values due
to an asymptote of TL seems to be reached (L∞is
similar to maximum TL). In this sense, the esti-
mated values of k(0.43 year-1) and t0(-0.36 years)
for males were higher than those previously calcu-
lated for individuals from Buenos Aires stock by
Barretto et al. (2011) (k =0.26 year-1 in spring and
0.21 year-1 in winter, t0 =-2.02 years in spring and
-2.90 years in winter). Since L∞and kare intimate-
ly related, the differences found in kfrom this
study could also be a consequence of females
reaching larger size in SMG, while males reached
smaller size as mentioned above. This caused a
decrease in the value of kfor females and an
increase in the value for males.
Taking into account the ages determined in our
study and the age at maturity (1.6 years for males
and 2.7 years for females) identified by
Rodrigues (2012), the differential growth for each
sex could be explained by the fact that once sex-
ual maturity is reached the growth rate decreased
(Brett 1979). In this way, females only begin to
use part of their energy for reproduction during
the second year of life and have a one-year
advantage over males to increase their size (Bar-
retto et al. 2011). Differences in growth could
also be determined by changes in environmental
conditions, water temperature and food availabil-
ity, which could affect metabolic rates (Dutil et al.
1999). The different growth rates may also be a
consequence of changes in the genetic character-
istics of the stocks (Renzi et al. 2009).
This work describes for the first time the growth
of P. brasiliensis in SMG and shows differences
with previous studies of this species from other
areas of the Argentine continental shelf. In agree-
ment with other authors (Braicovich and Timi
2008; Braicovich and Timi 2010; Rodrigues et al.
2010; López et al. 2012; Avigliano et al. 2015;
Braicovich et al. 2016), these preliminary results
provide data that suggest that the population of P.
brasiliensis from SMG constitutes a different
stock from those in CEB and the Argentine-
Uruguayan Common Fishing Zone (AUCFZ). In
this respect, defining stocks as discrete units is use-
ful for stock assessment and to propose manage-
ment measures (Ihssen et al. 1981). Future studies
on stock evaluation of P. brasiliensis in SMG
should contemplate appropriate methodologies for
data-poor species (Jiao et al. 2011). Determining
biological parameters that could limit and control
the management of fisheries is not an easy task,
however it is essential to understand as much as
possible about the resource in order to ensure that
its activity is sustainable in all aspects: the stock in
question, the community it inhabits and the society
that depends on the resource for its livelihood.
ACKNOWLEDGEMENTS
We thank the staff of Escuela Superior de
Ciencias Marinas (Universidad Nacional del
Comahue) and Instituto Nacional de Investiga-
ción y Desarrollo Pesquero (INIDEP) for provid-
ing the necessary tools to carry out this research.
110 MARINE AND FISHERY SCIENCES 32 (2): 103-114 (2019)
This work was partially funded by the Consejo
Interuniversitario Nacional (Argentina), based on
a grant awarded to the author of the work. We
thank CIMAS and Miguel Camarero “Kelo” for
assistance with sample collection. We also thank
Mariano Malvé, Damián Gil and Mauro Marcin-
kevicius for correcting the English language, and
making suggestions to improve the manuscript.
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114 MARINE AND FISHERY SCIENCES 32 (2): 103-114 (2019)
