JOURNAL OF CRUSTACEAN BIOLOGY, 33(2), 198-209, 2013
BY-CATCH ASSOCIATED WITH FISHERIES OF HETEROCARPUS VICARIUS (COSTA
RICA) AND HETEROCARPUS REEDI (CHILE) (DECAPODA: PANDALIDAE):
A SIX-YEAR STUDY (2004-2009)
Patricio M. Arana 1, Ingo S. Wehrtmann 2,3,∗, Juan Carlos Orellana 1,
Vanessa Nielsen-Muñoz 2,4, and Fresia Villalobos-Rojas 2
1 Escuela de Ciencias del Mar, Pontificia Universidad Católica de Valparaíso, P.O. Box 1020, Valparaíso, Chile
2 Unidad de Investigación Pesquera y Acuicultura (UNIP), Centro de Investigaciones en Ciencias del Mar y Limnología
(CIMAR), Ciudad de la Investigación, Universidad de Costa Rica, 11501-2060 San José, Costa Rica
3 Escuela de Biología, Universidad de Costa Rica, 11501-2060 San José, Costa Rica
4 Revista de Biología Tropical, Universidad de Costa Rica, 11501-2060 San José, Costa Rica
A B S T R A C T
Growing concern about the quantity and diversity of by-catch species caught in the bottom trawling nets of crustacean fisheries led us to
compare the quantity of by-catch recorded in Chilean and Costa Rican deep-water shrimp fisheries by year, latitude, and bathymetry. We
analyzed catches from 2143 trawl hauls between 2004 and 2009 from the fisheries of the northern nylon shrimp, Heterocarpus vicarius,
and the kolibri shrimp, Solenocera agassizii, off Costa Rica and the Chilean nylon shrimp, Heterocarpus reedi, off Chile. A catch index
was estimated to determine the quantity of by-catch retained for each kilogram of shrimp. The by-catch associated with the shrimp fisheries
of Costa Rica and Chile was mainly fishes and crustaceans; by-catch species diversity was considerably higher in Costa Rica compared to
that of Chile. In Chile, catches of H. reedi and by-catch were greater in the central and southern zone, whereas in Costa Rica, catches of
shrimp (H. vicarius, S. agassizii) and by-catch were higher in the central zone. In terms of bathymetry, the largest catches of shrimp and
by-catch came from the deep stratum (between 251 and 400 m) for both countries; the by-catch was noticeably larger in this stratum in
Costa Rica. The catch index revealed that for every 1 kg of shrimp caught in Costa Rica, 5.7 kg of by-catch were caught; in Chile, this ratio
was 1.1:1. However, the projected global by-catch was considerably higher for the fishery for H. reedi than for that of H. vicarius, which
is related to the landed volumes of the target species in Costa Rica and Chile. The encouraging results in the H. reedi-fishery concerning
technical measures to reduce by-catch and discards rates should be also considered for Central American deep-water shrimp fisheries.
KEY WORDS: by-catch, catch composition, deep-water shrimp, Heterocarpus spp., Solenocera agassizii
DOI: 10.1163/1937240X-00002123
INTRODUCTION
Benthic shrimp fisheries rely on trawl nets as the principal
fishing gear (Kelleher, 2005; Watson et al., 2006; Foster
and Vincent, 2010), and the use of trawls and dredges
has increased globally since the 1950s (Watson et al.,
2006). In general, bottom-trawling nets are considered to be
largely nonselective, catching and retaining large quantities
of species not targeted by the fishery; these species are
commonly referred to as “by-catch” (Broadhurst, 2000). By-
catch organisms are usually discarded due to the poor quality
of their meat, a lack of commercial interest, the fact that the
consumers are not accustomed to eating them, administrative
restrictions, and other social-economic reasons. However,
when returned to the sea, most of the by-catch has little or
no chance of survival (Villaseñor, 1997). Another relevant
aspect is that juveniles of the target resource may form
an important part of the by-catch, which can negatively
affect the commercial catches of the respective fisheries
(Broadhurst et al., 1996; Kennelly et al., 1998; Zhou, 2008;
Davies et al., 2009; Queirolo et al., 2011a). At present, the
∗ Corresponding author; e-mail: ingowehrtmann@gmx.de
information available on the quantity and quality of by-
catch extracted with bottom trawl nets is far from complete
(Kelleher, 2005; Davies et al., 2009). This gap is related
to the difficulty of verifying the information provided by
fishermen and to the fact that the participation of scientific
observers in commercial fishery operations is limited.
Interest in the by-catch associated with the extraction
of commercial species has grown in terms of its analysis
and evolution (Saila, 1983; Andrew and Pepperell, 1992;
Alverson et al., 1994; Kennelly, 1995; Harrington et al.,
2005; Zhou, 2008; Davies et al., 2009). Alverson et al.
(1994) defined by-catch as “discarded catch plus incidental
catch,” with incidental catch defined as the “retained catch
of non-targeted species.” In turn, Davies et al. (2009) defined
by-catch as B = Ct − Clm, where B is the total biomass of
the by-catch, Ct is the total biomass of the species caught in
the fishery haul, and Clm is the total catch landed or used.
In by-catch analyses, by-catch tends to be defined as the
percentage of undesired species with respect to the target
species of the fishery, as a ratio of one to the other, or
according to estimates of the total weight of the species that
© The Crustacean Society, 2013. Published by Brill NV, Leiden DOI:10.1163/1937240X-00002123
ARANA ET AL.: BY-CATCH ASSOCIATED WITH HETEROCARPUS SPP. FISHERIES 199
make up the by-catch in a given fishery, geographic region,
or globally (Alverson et al., 1994; Olsen et al., 2000; Davies
et al., 2009).
The existence of by-catch is a long-standing problem. The
negative effects created by discarding portions of fishery
catches have clearly led to a decline of marine ecosystems,
constituting a major world-wide problem (Watson et al.,
2006). Researchers are trying to evaluate the magnitude of
this matter and have proposed measures for its mitigation
and control (Graham, 1996; Hall et al., 2000; ECM, 2003;
Harrington et al., 2005; Krag et al., 2008; Matsuoka, 2008;
Davies et al., 2009). Specifically, trawling is considered
to be the most harmful fishery operation in terms of the
degradation of the marine floors, and this type of fishing is
thought to cause serious problems for fishery sustainability
(Broadhurst, 2000; Kelleher, 2005; Watson et al., 2006;
Foster and Vincent, 2010). Shrimp trawling, particularly in
the tropics, is considered to be the fishery that generates
the highest discard levels (Cook, 2001; Kelleher, 2005).
The ratio of by-catch to target resources in tropical and
subtropical shrimp fisheries often varies between 1:5 and
1:10, whereas in temperate and coldwater regions, this ratio
tends to be noticeably lower (Harris and Poiner, 1990;
Kelleher, 2005).
Most commercial fishery activities in Costa Rica are con-
centrated off the Pacific coast, where the main target species
of the deep-water fishery are the pandalid carideans Het-
erocarpus affinis Faxon, 1893 and Heterocarpus vicarius
Faxon, 1893, as well as Solenocera agassizii Faxon, 1893
(Decapoda: Penaeoidea: Solenoceridae). The latter species
is currently the most important in terms of annual shrimp
landings (Wehrtmann and Nielsen-Muñoz, 2009; Wehrt-
mann et al., 2009, 2012). The values of by-catch (discard) in
the Costa Rican shrimp fishery during 2000-2003 has been
estimated at 44% or 11 000 ton/year (Davies et al., 2009),
and the by-catch is generally 7.5 kg per 1 kg of shrimp (FAO,
2005).
In Chile, studies of the by-catch from trawl fishing
began in the early 1970s (Acuña et al., 2005). Information
on the resources caught with trawl nets used to extract
hake, Merluccius gayi (Guichenot, 1848), and the Chilean
nylon shrimp, Heterocarpus reedi Bahamonde, 1955, off
the central coast of the country (Yáñez, 1974; Yáñez and
Barbieri, 1974; Yáñez et al., 1974) revealed differences in
the spatial and seasonal abundance of the available species,
according to the kind of gear used. Qualitative descriptions
of the by-catch species of the H. reedi-fishery were provided
by other studies that focused on assessing the diversity of
deep-water fauna off central Chile (Acuña et al., 2005;
Orellana, 2006) and possibilities for reducing the by-catch
(Queirolo et al., 2011b).
Considering the discussions of and concerns with the sus-
tainability of deep-water fisheries and the impacts caused
by trawl fishing (Alverson et al., 1994; FAO, 2003; Althaus
et al., 2009), the present study analyzes the by-catch lev-
els associated with deep-water shrimp fisheries carried out
off Costa Rica (H. vicarius and S. agassizii) and Chile (H.
reedi). These two fisheries were selected, not only because
of their economical importance in Latin America, but princi-
pally because detailed by-catch data are available for a pro-
longed and comparable time period. Moreover, these fish-
eries refer to species of the same genus, distributed in differ-
ent geographic zones along the Pacific coast of Latin Amer-
ica, but in similar depth ranges. The principal objectives
of this work are: 1) to compare – over time, latitude and
bathymetry – the extraction of species associated with these
resources in two biogeographic regions, and 2) to determine
the catch index (CI). The results may contribute to our under-
standing of the ecological impacts of these deep-water fish-
eries and may facilitate the development of adequate man-
agement measures for each of the fisheries analyzed.
MATERIALS AND METHODS
This analysis of by-catch species relies on a historic data series compiled
during research projects carried out between 2004 and 2009. Data from
the Costa Rican fisheries of the northern nylon shrimp (H. vicarius) and
the kolibri shrimp (S. agassizii) were gathered as a part of a scientific
monitoring program of deep-water shrimp carried out off the Pacific coast
between 08°18′ and 10°12′N. The Chilean data were taken from assessment
cruises done on nylon shrimp (H. reedi) between 21°34′ and 38°29′S. In
both Costa Rica and Chile, the fishery operations were performed with trawl
nets with mesh sizes of 50 to 76 mm (2′′ and 3′′, respectively); no special
devices were employed to allow the escape or elimination of the by-catch.
The hauls lasted between 20 in Costa Rica and 30 min in Chile, and the
overall trawling depth range was from 102 to 653 m.
Data were collected by scientific personnel aboard commercial shrimp
vessels used for the research (length overall: 18-22 m). For each haul,
specially designed data sheets were completed with the fishery’s operational
data, setting and retrieval positions (using GPS readings), trawling depth
(using the vessel’s echo sounder), and the catch composition detailed by
species. The catch was weighed (recorded in kg) on board immediately after
retrieval. The by-catch species were quantified by separating them from the
target resource for each haul. The target catch (Costa Rica: H. vicarius, S.
agassizii; Chile: H. reedi) was placed in plastic bins arranged on the deck
of the vessel and later weighed and counted to determine the total catch in
weight.
The by-catch species were identified using the following literature: Costa
Rica: Bussing and López (1993), Garth (1958), Haig (1960), Hendrickx
and Salgado-Barragán (1991), Hendrickx (1995a, 1995b, 1995c, 1995d,
1997, 2000), Hendrickx and Estrada-Navarrete (1996), Keen (1971) and
Martin and Davis (2001); Chile: Pequeño (1994), Andrade and Báez (1980),
Retamal (1981) and Meléndez and Meneses (1989). Species that could not
be identified on board were separated and kept for identification on land by
specialists.
The databases generated in both countries were reviewed prior to
processing. For the general analysis of the information available, we used
only those hauls defined by the fishermen as being commercial catches, i.e.,
those in which >10 kg (Costa Rica) or >20 kg (Chile) of the target species
were caught. Later, for purposes of comparison, the records were grouped
by year, zone and depth. To determine possible associations of fauna for
H. vicarius, S. agassizii and H. reedi in terms of latitude, we grouped the
fishery hauls into three study sectors, i.e., the northern, central and southern
zones. The latitudinal limits for these sectors in both countries are presented
in Fig. 1. The records of both countries were also grouped into three depth
ranges: 100 to 250 m, 251 to 400 m, and 401 to 650 m; the latter depth
group refers only to Chile, because in Costa Rica all hauls analyzed were
carried out at depth above 400 m.
After grouping the data by zone and depth, the catch per unit effort
(CPUE) was calculated for the target species and the by-catch for each
of the two countries. Subsequently, we determined the arithmetical mean
values (x¯) with their standard deviations (s). Since all catch data showed
an asymmetric distribution, we applied the natural logarithm in order to
obtain a Gaussian distribution, which permitted to fulfill the assumption of
normality necessary to carry out the analysis of variance (Sokal and Rohlf,
1969).
To establish possible differences between CPUEs (of both target species
and by-catch) concerning zones, depth ranges and countries, the data were
analyzed by an analysis of variance (one factor ANOVA) with a significance
value of α = 0.05. When data did not comply with the homogeneity of
variance for the comparison of the means, a Kruskal-Wallis (KW) non-
parametric test was used.
200 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 33, NO. 2, 2013
Fig. 1. Identification of the study zones in Costa Rica and Chile. This figure is published in colour in the online edition of this journal, which can be
accessed via http://booksandjournals.brillonline.com/content/1937240x.
In case of significant differences between zones or depths, we applied
a posteriori multiple comparisons in order to determine which groups
presented differences. The Tukey test was used when variance homogeneity
was complied; if not, we utilized the Games-Howell test (Sokal and Rohlf,
1969). All analyses were carried out with the statistic program SPSS v.15.0.
In order to determine the amount of by-catch obtained with each fishery
haul per kg of deep-water shrimp, we calculated the catch index (CI) as
follows (Alverson et al., 1994):
CI =
∑
Catch of by catch (kg)
∑
Catch of objective shrimp (kg)
This index was calculated for different levels of shrimp catches in order
to elucidate how the by-catch varied according to greater or lower catches
of the target shrimps. Finally, the CI was used to project the catches of the
main species obtained as by-catch per country during the study period. In
Costa Rica, the global by-catch was estimated using annual landings of both
H. vicarius and S. agassizii. In Chile, this estimate was calculated in relation
to the annual global catch quotas of H. reedi.
RESULTS
The data bases compiled between 2004 and 2009 covered a
total of 2143 hauls (Table 1), from which we analyzed 1618
hauls with catches of the target resources, i.e., H. vicarius +
S. agassizii in Costa Rica and H. reedi in Chile. During this
time span, a total of 216 hauls were carried out in Costa Rica
(Fig. 2), 1402 were done in Chile (Fig. 3).
Catch Composition
The proportions of deep-water shrimp and by-catch differed
for the fisheries of the two countries studied. In Costa Rica,
the by-catch constituted between 82.3 and 96.8% of the total
annual catch in the period analyzed (Fig. 4). Moreover, the
by-catch increased as of 2007, coinciding with an important
reduction in the catch levels of H. vicarius and S. agassizii.
In Chile, the proportion of the by-catch ranged from 35.9%
(2006) to 57.1% (2008) (Fig. 4).
One important characteristic of the shrimp fisheries an-
alyzed from Costa Rica and Chile was the high diversity
of the by-catch. In general terms, the composition by taxo-
nomic group was similar in these two regions, dominated by
fishes, followed by crustaceans, mollusks and other groups
(Fig. 5). Species richness of the by-catch was considerably
higher in Costa Rica than in Chile; usually there were twice
the number of species per taxonomic group in Costa Rica
compared to those recorded in Chile, and the difference was
even more pronounced in mollusks (Costa Rica: 25 spp.;
Chile: 3 spp.). The catch in weight of the target species was
noticeably different between the two countries: on average,
the target species made up 9.2% of the total catch in Costa
Rica and 55.4% of the total catch in Chile (Table 2). In Costa
Rica, the by-catch of H. vicarius consisted mostly of diverse
crustacean species (62.1%), whereas in Chile, the most rel-
ARANA ET AL.: BY-CATCH ASSOCIATED WITH HETEROCARPUS SPP. FISHERIES 201
Table 1. Projects dealing with deep-water shrimp fisheries from 2004 to 2009, whose data were used for the present study: Heterocarpus vicarius and
Solenocera agassizii in Costa Rica and Heterocarpus reedi in Chile. ∗Number of haul with deep-water shrimp catches.
Year Name of the project Executing unit Study period Latitudinal Depth Number of
range range (m) hauls∗
2004 “Direct assessment of
nylon shrimp between
Regions II and VIII (FIP
2004-10)”
School of Marine
Sciences/The Pontifical
Catholic University of
Valparaíso, Chile
Jul 04-Sep 04 23°00′-37°00′ L.S. 120-548 251
“Development of standards
for a sustainable fishery of
the deep water shrimp
Heterocarpus vicarius (No.
111-A4-508)” 2004
Unit for Fishery Research
and Aquaculture (UNIP) of
the Research Center for
Marine Science and
Limnology (CIMAR),
University of Costa Rica
Jan 04-Dec 04 08°20′-10°12′ L.N. 210-347 37
2005 “Direct assessment of
nylon shrimp and prawn
between Regions II and
VIII (FIP 2005-08)”
School of Marine
Sciences/The Pontifical
Catholic University of
Valparaíso, Chile
Jul 05-Aug 05 24°30′-37°00′ L.S. 150-651 315
“Development of standards
for a sustainable fishery of
the deep water shrimp
Heterocarpus vicarius (No.
111-A4-508)” 2005
Unit for Fishery Research
and Aquaculture (UNIP) of
the Research Center for
Marine Science and
Limnology (CIMAR),
University of Costa Rica
Jan 05-Dec 05 08°46′-10°05′ L.N. 102-331 41
2006 “Direct assessment of
nylon shrimp between
Regions II and VIII (FIP
2006-11)”
Catholic University of the
North/University of
Concepción/Fisheries
Development Institute of
Chile (Instituto de Fomento
Pesquero, IFOP)
Oct 06-Dec 06 24°12′-36°42′ L.S. 154-653 232
“Development of standards
for a sustainable fishery of
the deep water shrimp
Heterocarpus vicarius (No.
111-A4-508)” 2006
Unit for Fishery Research
and Aquaculture (UNIP) of
the Research Center for
Marine Science and
Limnology (CIMAR),
University of Costa Rica
Jan 06-Dec 06 08°18′-09°31′ L.N. 172-329 29
2007 “Development of standards
for a sustainable fishery of
the deep water shrimp
Heterocarpus vicarius (No.
111-A4-508)” 2007
Unit for Fishery Research
and Aquaculture (UNIP) of
the Research Center for
Marine Science and
Limnology (CIMAR),
University of Costa Rica
Jan 07-Dec 07 08°18′-08°45′ L.N. 146-292 40
2008 “Direct assessment of
nylon shrimp between
Regions II and VIII (FIP
2008-17)”
Catholic University of the
North/University of
Concepción, Chile
Jan 08-Dec 08 25°16′-36°41′ L.S. 124-517 330
“Development of standards
for a sustainable fishery of
the deep water shrimp
Heterocarpus vicarius (No.
111-A4-508)” 2008
Unit for Fishery Research
and Aquaculture (UNIP) of
the Research Center for
Marine Science and
Limnology (CIMAR),
University of Costa Rica
Jan 08-Dec 08 08°19′-09°34′ L.N. 146-256 35
2009 “Direct assessment of
nylon shrimp between
Regions II and VIII (FIP
2009-16)”
Catholic University of the
North/University of
Concepción, Chile
Aug 09-Nov 09 25°08′-36°21′ L.S. 108-575 274
“Development of standards
for a sustainable fishery of
the deep water shrimp
Heterocarpus vicarius (No.
111-A4-508)” 2009
Unit for Fishery Research
and Aquaculture (UNIP) of
the Research Center for
Marine Science and
Limnology (CIMAR),
University of Costa Rica
Aug 09-Nov 09 09°15′-09°34′ L.N. 155-292 34
202 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 33, NO. 2, 2013
Fig. 2. Distribution of hauls carried out in the frame of a scientific monitoring program for deep-water shrimps (Heterocarpus vicarius and Solenocera
agassizii) along the Pacific coast of Costa Rica (2004-2009). This figure is published in colour in the online edition of this journal, which can be accessed
via http://booksandjournals.brillonline.com/content/1937240x.
evant by-catch groups were crustaceans (22.9%) and fishes
(21.1%) (Table 2).
In Costa Rica, the most common crustaceans were Squilla
biformis Bigelow, 1891 (12.1%) and Pleuroncodes monodon
(H. Milne Edwards, 1837) (12.5%). Although no detailed
identification of the fish species present in each haul was
recorded during 2004-2009, this group made up on average
28.4% of the total weight caught in the study period
(Table 2). In Chile, the crustaceans that contributed the most
in weight were the squat lobsters Cervimunida johni Porter,
1903 (11.0%) and Pleuroncodes monodon (3.4%). In the
fish group, the main resources associated with the catches
of H. reedi were South Pacific hake (Merluccius gayi), rat-
tail (Caelorhynchus fasciatus (Günther, 1878)) and bigeye
Fig. 3. Distribution of hauls along the Pacific coast of Chile carried out as part of assessment cruises for Heterocarpus reedi (2004-2009). This figure is
published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/1937240x.
ARANA ET AL.: BY-CATCH ASSOCIATED WITH HETEROCARPUS SPP. FISHERIES 203
Fig. 4. Comparison of mean by-catch per year (expressed as percent of
total catch) recorded during research projects in Chile and Costa Rica
(2004-2009). n.d. = no data available.
Fig. 5. Number of species per taxonomic group in the by-catch of deep-
water shrimps in Costa Rica and Chile (2004-2009).
Table 2. Composition (by percentage) of the catch in weight (kg) for
taxonomic groups in Costa Rica and Chile (2004-2009).
Year Shrimp Fishes Crustaceans Mollusks Others Total
catch
(kg)
Costa Rica
2004 12.1 14.0 73.8 0.0 0.0 9919
2005 17.4 27.0 55.4 0.1 0.1 7640
2006 12.7 35.2 51.9 0.1 0.0 2132
2007 2.7 12.6 84.6 0.1 0.0 12 233
2008 3.7 36.2 59.1 0.6 0.4 6629
2009 4.2 44.4 51.1 0.2 0.1 8148
Total (%) 9.2 28.4 62.1 0.2 0.1
Chile
2004 49.9 38.1 10.2 1.7 0.1 64 898
2005 53.3 35.2 10.7 0.5 0.3 120 271
2006 64.1 16.2 19.3 0.5 0.0 65 701
2007 0
2008 42.9 10.6 45.8 0.7 0.0 163 360
2009 69.1 16.1 14.8 0.1 0.0 150 867
Total (%) 55.4 21.1 22.9 0.6 0.1
Fig. 6. Average deep-water shrimp catches and by-catch (expressed as
percent of total catches), by zone (2004-2009).
flounder (Hippoglossina macrops Steindachner, 1876) with
9.1%, 3.5% and 3.2%, respectively.
Analysis of CPUE of Target Species and By-catch
In Costa Rica, the central zone accounted for more than 60%
of all the by-catch obtained during the study period in all
three zones (Fig. 6). Considering the low number of hauls
carried out in the northern Pacific zone of Costa Rica, only
the central and southern zones were compared statistically:
we did not detect significant differences regarding the
CPUEs of neither the two target species (ANOVA; F =
0.159; p = 0.690) nor the by-catch (ANOVA; F = 0.317;
p = 0.574). In Chile, the highest percentage of total
catches of H. reedi was recorded for the southern zone
(Fig. 6); however, the highest average CPUE (682.5 kg/h)
was obtained in the southern zone of Costa Rica (Tables 3
and 4). The statistical analysis revealed that the CPUEs of
both the target species H. reedi (ANOVA; F = 40.64;
p = 0.000) and the by-catch (KW; χ2 = 39.76; p = 0.000)
increased significantly from north to south. Moreover, and in
contrast to the situation in Costa Rica, landings of the target
species (H. reedi) were always higher than that of the by-
catch.
The catch analyses of the two target species in Costa Rica
per depth range revealed that the extracted average volumes
between 100-250 and 251-400 m were similar; no hauls were
carried out deeper than 400 m (Fig. 7). By-catch was more
abundant in the depth range of 100-250 m. In Chile, highest
values for H. reedi and by-catch were recorded between 251-
400 m, followed by the depth range of 100-250 m (Fig. 7). In
Costa Rica CPUEs values of the target species were higher
204 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 33, NO. 2, 2013
Table 3. Deep-water shrimp CPUE (kg/h) values per zone, depth and
country, n = number of hauls (2004-2009).
Deep-water
shrimp
Costa Rica Chile
n Media Standard n Media Standard
(kg/h) deviation (kg/h) deviation
Zone
Northern 6 19.6 16.9 256 327.6 304.0
Central 169 53.4 87.8 722 505.7 442.0
Southern 26 77.2 132.1 422 682.5 635.8
Depth (m)
100-250 112 40.1 71.1 251 648.1 662.9
251-400 89 74.8 113.4 985 541.0 473.8
401-650 – – – 164 252.9 227.3
in 251-400 m (74.8 kg/h), whereas CPUEs values for by-
catch were higher in 100-250 m (604.5 kg/h) (Tables 3 and
4). However, no statistically significant differences of the
CPUEs among the depth ranges were recorded neither for
the target species (KW; χ2 = 1.649; p = 0.199) nor for the
by-catch (ANOVA; F = 2.923; p = 0.089).
In Chile, CPUEs of both the target species and the by-
catch decreased with depth, with the highest catch rates in
the depth range of 100-250 m. CPUEs of both the target
species (KW; χ2 = 102.93; p = 0.000) and its by-catch
(ANOVA; F = 27.11; p = 0.000) were significantly
different between depth ranges. Further statistical analysis
provided evidence that there were no significant differences
of the CPUEs comparing the ranges of 100-250 m and 251-
400 m neither for H. reedi (Games-Howell; p = 0.381)
nor for the by-catch (Tukey; p = 0.331). However, CPUEs
of the target species (Games-Howell; p = 0.000) and the
by-catch (Tukey; p = 0.000) were statistically different at
depth > 400.
Overall average CPUEs for the target species was 55.4 and
526.4 kg/h in Costa Rica and Chile, respectively; the values
for both countries were statistically significant different
(KW; χ2 = 448.73; p = 0.000). In the case of the by-catch
(Fig. 8), the mean CPUE for Costa Rica (598.0 kg/h) was
significantly higher (KW; χ2 = 47.37; p = 0.000) than the
same value for Chile (340.8 kg/h).
Table 4. By-catch CPUE (kg/h) values per zone, depth and country, n =
number of hauls (2004-2009).
By-catch Costa Rica Chile
n Media Standard n Media Standard
(kg/h) deviation (kg/h) deviation
Zone
Northern 6 956.6 726.4 256 288.6 477.3
Central 169 543.4 694.9 722 321.1 482.0
Southern 26 870.0 1830.4 422 406.2 449.4
Depth (m)
100-250 112 604.5 793.0 251 474.9 624.3
251-400 89 589.9 1070.8 985 335.7 449.3
401-650 – – – 164 166.0 217.8
Fig. 7. Average deep-water shrimp catches and by-catch (expressed as
percent of total catches), by depth range (2004-2009). n.d. = no data
available.
Catch Index (CI)
The catch index (CI) revealed that the quantity of by-catch
in weight obtained for each kg of shrimp was noticeably
greater in Costa Rica than in Chile (Fig. 9). A comparison
of catches of the target species with those of the by-catch
revealed in both countries a close relationship between high
by-catch levels and low catches of the target species; this
situation changed when the retention of the desired species
incremented (Fig. 10, Table 5).
In Costa Rica, considering for example catches greater
than 10 kg of the target species (H. vicarius and S. agas-
sizii), for each kg of shrimp an average of 5.7 kg of by-catch
organisms were caught (CI = 1:5.7). In Chile, when consid-
ering hauls with catches of H. reedi greater than 20 kg, for
Fig. 8. CPUE values for deep-water shrimp catches and by-catch per
country (Costa Rica, Chile: 2004-2009).
ARANA ET AL.: BY-CATCH ASSOCIATED WITH HETEROCARPUS SPP. FISHERIES 205
Fig. 9. Average annual catch index determined for the Heterocarpus
vicarius and Solenocera agassizii (Costa Rica) and H. reedi (Chile) fisheries
(2004-2009).
Fig. 10. Average estimated catch indexes for different minimum catch
amounts of deep-water shrimps in Costa Rica (Heterocarpus vicarius and
Solenocera agassizii) and Chile (Heterocarpus reedi) during the study
period 2004-2009. Numbers over bars indicate CI values.
Table 5. Catch indexes obtained for the Heterocarpus vicarius and
Solenocera agassizii fisheries of Costa Rica and the H. reedi fishery of Chile
(2004-2009). n = number of hauls.
Year Catch index according to different
levels (kg) of captured shrimps
>30 >20 >15 >10 >9 >8 >7 >6 >5
Costa Rica
2004 1.9 2.8 2.7 3.7 5.0 5.9 5.8 15.2 17.0
2005 2.6 2.4 3.3 4.8 4.8 4.8 5.0 7.8 7.9
2006 1.0 1.4 1.6 3.0 3.0 3.3 3.3 5.9 5.9
2007 4.2 4.5 4.0 3.8 5.2 5.2 5.2 27.0 27.0
2008 7.7 5.9 5.7 6.1 6.1 6.3 9.6 9.6 11.4
2009 12.3 12.6 15.4 15.1 15.1 15.1 17.7 18.0 22.3
n 35 47 70 91 95 100 107 114 123
Chile
2004 1.1 1.1 1.3 1.8 1.9 2.1 2.1 2.3 2.3
2005 1.0 1.1 1.3 1.7 1.9 2.1 2.1 2.2 2.2
2006 0.9 1.1 1.2 1.4 1.4 1.3 1.5 1.5 1.5
2007 – – – – – – – – –
2008 1.0 1.0 2.7 3.8 4.0 4.0 4.4 4.4 4.3
2009 1.0 1.1 1.6 2.6 2.7 2.7 2.7 2.7 3.0
n 1344 1402 1473 1531 1544 1567 1576 1582 1589
Fig. 11. Estimated global annual by-catch for the Heterocarpus vicarius
and Solenocera agassizii (Costa Rica) and Heterocarpus reedi (Chile)
fisheries (2004-2009). n.d. = no data.
each kg of shrimp, 1.1 kg of by-catch was recorded (CI =
1:1.1). In Costa Rica, the CI incremented notoriously as of
2008, reaching a ratio of 1:15.1 in 2009 (Table 5).
According to the average catch indexes (CI) of each
country and the annual landings recorded for Costa Rica
(2004-2007) and Chile (2004-2009), the annual reported by-
catches for the H. vicarius and S. agassizii fisheries were
1041 to 3147 ton and for the H. reedi fishery were 4103
to 4476 ton (Fig. 11). The amount of by-catch extracted
annually in Chile showed a certain degree of stability (4058
to 4476 ton) and was associated with similar landings (3644
to 3956 ton) between years. In Costa Rica, however, the by-
catch value peaked in 2005, followed by a sustained drop
in the total amounts that was directly associated with the
decline of the H. vicarius fishery following that year.
DISCUSSION
The proportion of by-catch in Costa Rica for each of
the years analyzed was notoriously greater than the target
resource catches, with values of up to 96.8% (mean 90.8%)
of the total catch (Fig. 4). Likewise, Sarmiento-Náfate et
al. (2007) found that the by-catch of the shrimp fisheries in
Mexico can reach up to 96.7% of the total catch. On the
contrary, for the Chilean trawl fishery of H. reedi, Melo et
al. (2006) estimated that the catches consisted mainly of the
resource target, with only 35% being by-catch; a similar
value was found in the present research. In general, and
according to various direct assessments carried out for H.
206 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 33, NO. 2, 2013
reedi (Arana et al., 2004, 2006; Acuña et al., 2007, 2009,
2010), it seems safe to assume that in Chile, the by-catch
associated with the H. reedi-fishery is generally less than
50% of the total catch measured in weight. The observed
differences of by-catch rates between the Costa Rican and
Chilean deep-water shrimp fisheries are in agreement with
the conclusion of Gillett (2008) that in terms of the quantity
of by-catch extracted, problems are less severe in the cold-
water trawl fisheries as opposed to those in tropical waters.
The by-catch in the Costa Rican deep-water shrimp fish-
ery was highly diverse when compared to that of the H.
reedi-fishery in Chile. This pattern is not surprising, con-
sidering that Costa Rica is known for its high biodiversity,
including the marine realm (Wehrtmann and Cortés, 2009).
The catches in both countries contained relatively similar
percentages in terms of the number of species per taxo-
nomic groups (Fig. 5). However, we observed substantial
weight differences between these groups: crustaceans pre-
dominated (62.1%) in the H. vicarius fishery, whereas sim-
ilar percentages of fishes (21.1%) and crustaceans (22.9%)
were found in the H. reedi-fishery (Table 2). On the other
hand, at species level, the composition of fauna was different
in these two regions. In Costa Rica, the crustacean by-catch
was mainly made up of Squilla biformis, Plesionika trispinus
Squirres and Barragan, 1976, and Pleuroncodes monodon
(Wehrtmann and Echeverría-Sáenz, 2007). In Chile, how-
ever, the by-catch of the nylon shrimp fishery was made up
of several fish species, being the South Pacific hake (Merluc-
cius gayi) the most important one (Melo et al., 2003; Acuña
et al., 2005; Orellana, 2006; Queirolo et al., 2011b).
The composition of the by-catch observed in the H.
vicarius-fishery coincided with that reported by Puentes et
al. (2007) for the fisheries for Solenocera agassizii and
Farfantepenaeus spp. carried out in the Colombian Pacific,
where the by-catch was dominated mainly by 54 fish and 11
crustacean species. It should also be noted that, in Colombia,
mollusks, and other benthic taxonomic groups were poorly
represented. However, this might be a result of the relatively
short study period (one year), and it might be speculated
that not all by-catch species were separated and identified,
because Puentes et al. (2007) mentioned as mollusk just one
squid species, not even one bivalve or gastropod species.
According to Hall et al. (2000), tropical shrimp fisheries
present average CI values from 1:3 to 1:15. Other studies
(Harris and Poiner, 1990; Andrew and Pepperell, 1992;
Alverson et al., 1994; Ambrose et al., 2005) have reported
this index to be 1:5 in temperate zones and 1:10 in tropical
zones, depending on the availability and reliability of the
data generated in each of the fisheries analyzed. The CI
estimated for Costa Rica fell within the ranges indicated
by the aforementioned authors, averaging 5.7 kg of by-
catch for each kg of shrimp caught (1:5.7). The CI in Chile,
considering catches of H. reedi in hauls with over 20 kg of
the target species, however, was 1:1.1, which was lower than
indicated by earlier estimates for temperate waters (Harris
and Poiner, 1990; Kelleher, 2005).
In Costa Rica, practically all of the by-catch obtained
in the deep-water shrimp fishery is discarded, even those
species that have commercial value in other Central Ameri-
can countries (Wehrtmann and Nielsen-Muñoz, 2009). Sar-
miento-Náfate et al. (2007) found a similar situation for
the Gulf of Tehuantepec, Mexico, where only 10% of the
by-catch was used for commercial purposes. According to
Kelleher (2005), one of the main problems associated with
discards is the use of small vessels with limited space for
storing the catches. Other problematic factors include the
difficulty of conservation and low prices at local markets.
In contrast, an important fraction of the by-catch from the
H. reedi-fishery is not discarded, as it contains species of
commercial interest. Thus, landings of South Pacific hake
and squat lobsters are authorized in quantities that do not
exceed (for each of these) 10% of the total catch in weight
of the target resource (Ley General de Pesca y Acuicultura,
Chile, 1992). Catches of these species were considered to be
by-catch in the present research. If the portion of by-catch
that is of commercial interest is taken in account, the by-
catch ratio or CI will decrease at least by 9.1%, from 1:1.1
to 1:1.0.
On the other hand, it should be taken in consideration that
the by-catch in both countries was quantified by using data
from fishery hauls with and also without the presence of the
resource target obtained in direct assessment projects. This
generated an overestimation of the by-catch values obtained
for each kg of shrimp caught and, therefore, an erroneous
perception of the real levels of by-catch extracted. Hence,
it is likely that the values indicated by Oceana (2005) for
Chile (81.2%) were overestimates, as they considered the
entirety of the hauls carried out during assessment projects
of crustaceans, and did not represent the actual catches of
the commercial fleet.
The results of the present research revealed differences
in the quantity of by-catch according to the fishery zone,
the depth at which the extractive operations were carried
out (Table 4), and catch concentrations of the target species
(Fig. 9, Table 5). These findings suggest that the density of
the by-catch could be relatively constant over the trawled
bottoms, whereas the shrimps are concentrated in certain
zones and depths, which results in different indexes found in
relation to the variation in the catches of the target species.
This situation shows that the data used to calculate the by-
catch must be considered carefully since the use of different
minima of catch per haul of the target species can distort this
index noticeably.
We assume that the captains dedicated to crustacean
fisheries know the zones and depths in which the foci
of abundance of the target resource are found, leading to
high catches and relatively few by-catch in these areas.
Nevertheless, it is also likely that the proportion of the by-
catch increases each time that the captain perform fishery
hauls beyond the normal aggregation sites of the target
resource or when occasionally prospecting on bottoms in
search of new fishing spots.
The projected global by-catch was considerably higher
for the H. reedi-fishery than for the H. vicarius-fishery
(Fig. 11). This apparent contradiction was related to the
landed volume of each of these species: although in Costa
Rica the percentages of by-catch were higher than those
recorded in Chile, the landings between 2004 and 2006 did
not exceed 672 ton, with yearly by-catch figures ranging
between 2039 ton and 3147 ton. Furthermore, in 2007, the
ARANA ET AL.: BY-CATCH ASSOCIATED WITH HETEROCARPUS SPP. FISHERIES 207
by-catch dropped (1041 ton) along with the reduction of the
target catch (278 ton). In turn, in Chile, the landings of the
industrial sector remained stable, fluctuating between 3644
and 3956 ton (2004-2009) and the projected by-catch was
between 4187 and 4354 tons per year.
Several authors have tried to determine the total by-catch
at a global level. The estimates exceeded 38.5 million tons,
or 40.4% of the global landings of marine resources (Davies
et al., 2009), which represents hypothetical economic losses
near one billion dollars annually (Cook, 2001; FAO, 2003).
Thus, it is urgently important to adopt measures to reduce
the by-catch and discard rates (Harrington et al., 2005; Zhou,
2008). The elimination of resources of low economic value
and/or that do not comply with the fishery regulations (e.g.
below the minimum size) constitutes mortalities that are not
taken into consideration in the fishery statistics, leading to
biased estimations of the biomass of exploited resources.
At the same time, this situation impedes determining the
mortality levels to which the non-target species of the
fishery are subjected (Alverson et al., 1994; Kennelly, 1995;
Alverson and Hughes, 1996; Hall et al., 2000; Zhou, 2008).
The present study shows high volumes of by-catch result-
ing from the fisheries for H. vicarius and H. reedi. This by-
catch should be reduced in order to conserve the extracted
resources and protect the marine environment. Several mit-
igation measures have been suggested for the shrimp trawl
fishery, but these only achieve partial results since they do
not restrict the use of trawl nets as part of the extractive ac-
tivities. The most recent research on the by-catch was aimed
at determining how to reduce the catch of these organisms,
carrying out technological modifications of the nets in or-
der to increase selectivity (Broadhurst, 2000; ECM, 2000;
Queirolo et al., 2011a) and incorporating the use of escape
devices to diminish the retention of undesired species (Ken-
nelly, 2007). Other indirect actions should aim at using the
presently discarded and unused species and assigning eco-
nomic incentives to businesses and fishermen in order to in-
volve them in the development of ideas and projects intended
to decrease the by-catch in their extractive operations (Cerda
et al., 2009). Unlike the Costa Rican case, in Chile manage-
ment measures include prohibitions on landings of by-catch
species of other commercial species and on returning dis-
carded specimens to the sea. Although these measures have
contributed partially to the desired objective, the fishermen
do not always comply with them, despite being subjected to
substantial fines. Therefore, greater attention has been paid
to the search for technological solutions. In the case of the
nylon shrimp fishery, the use of square meshes of 56 mm and
the use of grids (by-catch reduction devices) to separate the
by-catch from the target catch have reduced the by-catch by
around 40%, with a loss of the target catch of around 10%
(Queirolo et al., 2011a). Considering that shrimp trawl fish-
eries, and tropical shrimp fisheries in particular, are the sin-
gle greatest source of discards (Kelleher, 2005), we strongly
support additional studies aimed to reduce the by-catch and
discard rates in tropical shrimp fisheries. The encouraging
results in the H. reedi-fishery (Queirolo et al., 2011a) should
be used to introduce these technical measures also in Central
American deep-water shrimp fisheries.
ACKNOWLEDGEMENTS
The authors (PA and JCO) thank the Fondo de Investigación Pesquera (FIP)
for funding the fishery assessment projects carried out off the central coast
of Chile and for the availability of the data bases used to develop the
present project. The researchers from Costa Rica (ISW, VNM and FVR)
are thankful to the company The Rainbow Jewels, S.A., Puntarenas, for
their ongoing support for monitoring the deepwater resources of Pacific
Costa Rica. Likewise, thanks a lot to the captains (Rigo and Esteban:
“Mecate”), crews of the shrimp trawlers ONUVA and SULTANA, and all
the students who helped us with the collection and handling of the samples.
Financial support was made available from the German company Ristic
AG, Oberferrieden, the Costa Rican company The Rainbow Jewels S.A.,
Puntarenas, and the University of Costa Rica (projects V.I. No. 111-A4-
508, V.I. No. 808-A9-536 and V.I. No. 808-A9-537). Additional funding
was provided by the “Program University – Enterprise for Sustainable
Development” (PUEDES) of the Council of Central American Universities
(CSUCA), the German Society for Technical Cooperation (GTZ) and
the University of Kassel, Germany. Furthermore, we are grateful for the
logistic support provided by the Center of Marine Research and Limnology
(CIMAR) and the School of Biology, University of Costa Rica. We are
grateful to Ing. Álvaro De Caso for his collaboration with the statistical
analysis of the fishery data. Danielle Barriga and Tayler Clarke revised and
improved the English, which is greatly appreciated. Finally, we are thankful
to Raquel Romero-Emilia, who helped us to prepare the final version of the
figures.
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