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Animal Conservation. Print ISSN 1367-9430

Using distribution patterns of small fishes to assess small

fish by-catch in tropical shrimp trawl fisheries
S. J. Foster1 & F. Arreguin-Sánchez2
1 Fisheries Centre, The University of British Columbia, Vancouver, British Columbia, Canada
2 Centro Interdisciplinario de Ciencias Marinas del Instituto Politécnico Nacional – CICIMAR-IPN, La Paz, Baja California Sur, México

Keywords
protected areas; fisheries; impact;
non-target species; conservation.
Correspondence
Sarah Foster, Project Seahorse, Fisheries
Centre, The University of British Columbia,
2202 Main Mall, Vancouver, British
Columbia V6T 1Z4, Canada. Tel:
+604 827 5139
Email: s.foster@fisheries.ubc.ca
Editor: Iain Gordon
Received 24 April 2012; accepted 23
August 2013
doi:10.1111/acv.12078
Abstract
Ecologically sound fisheries management and improving future food security
require that small fish by-catch in tropical shrimp trawl fisheries is maintained at,
or reduced to, sustainable levels; restricting trawling in particular places, or at
particular times, has been suggested as a means for achieving this goal. The
purpose of our research was to compare patterns in occurrence, density and body
size across depth, latitude and time for four small fish taxa caught as by-catch in
the southern Gulf of California shrimp trawl fishery: Diplectrum spp., Prionotus
stephanophrys, Pseudupeneus grandisquamis and Stellifer illecebrosus. We then
used these results to explore the potential for trawler impacts on these taxa, and
the possible placement and timing of fishing restrictions to mitigate potential
impacts. Our results confirmed, however, the difficulties of regulating such fish-
eries for multiple by-catch species, in that their distribution patterns varied in a
way that precludes a ‘one size fits all’ solution. The four taxa analysed – only four
of the hundreds obtained as by-catch in this fishery – exhibited distribution
patterns at odds with one another. Observed intra- and inter-taxon variations in
the relative importance of different spatial and temporal variables in determining
occurrence, density and size argues that several permanent trawl closures covering
a range of depths and latitudes, and not temporal ones, might be required to
mitigate potential trawl impacts on these fishes. Our results also suggested a higher
potential for impact on S. illecebrosus than the other taxa: occurrence and density
of the former declined, whereas occurrence or density of the others increased
across the study area as the fishing season progressed.

Introduction mitigating the effects of trawling has been mostly focused in

Commitments to ecologically sound fisheries management
(e.g. Sherman et al., 2005), and the importance of fisheries
to food security (e.g. Delgado et al., 2003), require that
small fish by-catch is maintained at, or reduced to, sustain-
able levels. By-catch is the non-target part of the catch and
is either discarded at sea or retained for human or animal
consumption (Andrew & Pepperell, 1992). Previous work
has shown that species taken as by-catch are sometimes
harvested at unsustainable levels (even in regulated fisher-
ies), threatening species diversity and ecosystem health
(Hall, 1996). Shrimp trawlers pose a particular challenge in
that they catch many million tonnes of non-target species
each year (Alverson et al., 1994; Kelleher, 2004), with tropi-
cal shrimp fisheries having the highest discard rates of all
fisheries (Kelleher, 2004). The majority of by-catch species
in tropical shrimp fisheries are comprised of fishes that
mature at less than 20 cm (Alverson et al., 1994), hereafter
collectively referred to as ‘small fishes’. Yet, research into
temperate or developed tropical systems, and on charis-
matic or commercially important species. We therefore
know remarkably little about how shrimp trawling practices
affect the majority of the species at the bottom of the nets.
Although technological advances in the shrimp trawl
industry have helped reduce by-catch of some groups of
fauna, by-catch of small fish species is still a problem, pri-
marily because they are the same size as tropical shrimps
(Clucas, 1997; Cochrane, 2002). Most by-catch reduction
devices (BRDs) exclude individuals that are larger than
target shrimps (Broadhurst, 2000; Eayrs, 2007). For
example, when properly implemented, turtle excluder
devices significantly reduce by-catch of turtles and other
large fishes, and implementation of BRDs in US shrimp
fisheries have reduced by-catch of commercially important
red snapper (as reviewed in Broadhurst, 2000 and Eayrs,
2007). Ongoing work continues to explore the use of juvenile
and trash excluder devices – rigid grid sorting devices
installed in the part of the trawl where fishes are retained –

Animal Conservation •• (2013) ••–•• © 2013 The Zoological Society of London 1

Mitigating small fish by-catch based on distribution patterns
to allow smaller fish and shrimps to escape, but high con-
struction costs may limit implementation (Eayrs, 2007). In
spite of technological advances, by-catch levels are still high;
discard rates for all taxa averaged 56% across tropical
shrimp fisheries (Kelleher, 2004). Hence, it may never be
possible to find a technological solution that works for all
small fishes, because of their diverse behaviour patterns,
physiological conditions and morphologies (Bublitz, 1995;
Loverich, 1995).
Closing a shrimp fishery in particular areas or at par-
ticular times (or a combination thereof), thereby reducing
trawl intensity in space or time, may be the most likely
means to adjust by-catch of the hundreds of small fish
species caught by tropical shrimp trawlers. Managed access
or closures may be useful for protecting a variety of fishes
with divergent life-history strategies (Winemiller & Rose,
1992). Trawling restrictions not only reduce incidental
fishing mortality but also reduce habitat damage, thereby
addressing both direct and indirect impacts on by-catch
species (Kaiser, 2000; Stefansson & Rosenberg, 2005).
Restricting any closures to periods of high fish abundance
or when fishes are most vulnerable to overexploitation (e.g.
during spawning aggregations), but allowing fishing to
occur the remainder of the season, may reduce economic
costs of the closures and so ease their implementation
(Andrew & Pepperell, 1992). What is not clear is how such
fishery closures should be applied to address the by-catch
of small fishes – whether potential impacts on these taxa
could be mitigated with a common closure plan, or if each
fish species requires its own closure placed uniquely in
space and time.
Information on species distribution is useful to assess the
potential value of spatio-temporal restrictions in mitigating
fisheries’ impacts (Morgan & Chuenpagdee, 2003). For
example, one could map spatio-temporal patterns of fishing
effort on patterns of species densities to locate ‘by-catch
hotspots’, regions of greatest overlap and thus locations
where managed areas would have greatest potential for
impact. Similarly, information on size distributions in space
and time could inform the location of spawning grounds or
periods of high vulnerability, such as reproductive peaks.
Unfortunately, few such data are available for the small
fishes caught as by-catch in tropical shrimp trawls (e.g.
Stobutzki, Miller & Brewer, 2001).
Thus, the purpose of our research was to test the hypoth-
esis that four small fish taxa caught as by-catch in a tropical
shrimp trawl fishery shared common patterns of occurrence,
density, and body size across space (depth, latitude) and
time (day). We then used the results to explore suggestions
of (1) potential for trawler impact on these taxa, and (2) the
placement and timing of fishing restrictions to mitigate
potential impacts. Ultimately, we wanted to see if there was
a commonality in distribution patterns across taxa that
could inform where and when to implement fishing
restrictions, if one wanted them. The research was focused
in the southern Gulf of California (Mexico) and on four
small fish taxa: Diplectrum spp., Prionotus stephanophrys,
Pseudupeneus grandisquamis and Stellifer illecebrosus.
2 Animal Conservation •• (2013) ••–•• © 2013 The Zoological Society of London
S. J. Foster and F. Arreguin-Sánchez
USA
N
Mexico
Sinaloa
Mazatlán Gulf of california
50 km
Gulf of california Nayarit
Tepic
a. b.
c. d.
Figure 1 Map of Gulf of California, México, indicating sampled area
(inset), station locations and case study taxa: (a) Diplectrum labarum,
(b) Pseudupeneus grandisquamis, (c) Prionotus stephanophrys and
(d) Stellifer illecebrosus. Species photos © CICIMAR.
Materials and methods
Study site
Our work focused on the southern Gulf of California, one
of the gulf’s three biogeographic regions (Walker, 1960;
Fig. 1). More than 1000 large industrial shrimp trawlers
sweep an area of the sea floor equivalent to two times that of
the Gulf on an annual basis (Brusca et al., 2005), inciden-
tally catching more than 100 species of fish (Perez-Mellado
& Findley, 1985). A decrease in diversity and biomass of
by-catch species over the last 50 years has been inferred, but
not confirmed, from limited scientific and anecdotal infor-
mation (Brusca et al., 2005; Sáenz-Arroyo et al., 2005;
Lozano-Montes, Pitcher & Haggen, 2008).
Study species
We chose four commonly caught small fish taxa in the
Gulf’s trawl fishery as case studies for this research:
Diplectrum spp. (sandperch; samples were erroneously con-
sidered to consist of a single species at time of data collec-
tion, but in fact contained individuals of Diplectrum
labarum and D. pacificum at least), lumptail searobin
Pr. stephanophrys, bigscale goatfish Ps. grandisquamis, and
silver stardrum S. illecebrosus (Fig. 1). We chose these taxa
because they were good representatives to determine if a
‘one size fits all’ approach to fishing restrictions is tenable.
They all had potentially high abundance in the by-catch (to
ensure sufficient sample sizes), were thought to spend their
entire life cycle on the fishing grounds (V. Cruz, CICIMAR,
pers. comm.), and each belonged to one of the more com-
S. J. Foster and F. Arreguin-Sánchez
monly caught families in this fishery (Manjarrez Acosta,
2001; CICIMAR, unpubl. data). All are small in size
(maximum sampled total length ranged from 19 to 31 cm
across taxa; Foster & Vincent, 2010) and associated with
benthic habitats (sandy and muddy bottoms; Eschmeyer,
Herald & Hammann, 1983; Chao, 1995; Heemstra, 1995;
Schneider, 1995). A companion paper presents details on
survival, growth and reproduction of the focal taxa (Foster
& Vincent, 2010). All taxa exhibited prolonged periods of
reproduction in the southern Gulf of California, with
mature individuals found throughout the trawl season (Sep-
tember to March) (Foster & Vincent, 2010).
Data collection
We participated in fishing trips on two industrial shrimp
trawlers during the 2006–2007 fishing season to sample the
focal taxa in the by-catch. We collected information on the
location and date of capture, as well as the quantity and size
of the taxa when caught by the trawlers. Samples of all taxa
were collected and processed at 38 fixed stations across the
shrimp fishing grounds for the entire 2006–2007 fishing
season (Fig. 1). The stations were the same as those sampled
annually by Mexican fisheries authorities to assess shrimp
population status and determine the opening dates for the
fishery. They were spread across an area measuring
11 600 km2 and so covered most of the southern Gulf fishing
grounds, and ranged in depth from 7 to 62 m. The industrial
shrimp fishery in the gulf operates from September to
March. Sampling details are described in Foster & Vincent
(2010). The vessels used for sampling were typical of the
industrial shrimp trawlers used throughout the Gulf (see
Gillett, 2008), deploying paired otter trawls with mesh size
of 4.5–5.0 cm.
Response variables of interest for all taxa were occur-
rence (presence/absence), density and total length (LT, mm).
Density was used instead of counts to account for variation
in gear (net length) and effort (tow duration) during sam-
pling. Density was calculated as the total number of indi-
viduals per kilometre2 per hour of trawling (n km−2 h−1). The
Gulf’s trawlers fish with two nets, so area swept by the
gear ≈ 2 × width of the net opening × distance trawled,
where width of the net opening ≈ net length × 0.55 (J. T.
Nieto, Escuela Nacional de Ingeniería Pesquera, Nayarit,
pers. comm.). The distance trawled was obtained using a
Global Positioning System (GPS).
Predictor variables included depth, latitude and time.
Understanding how a species is distributed across latitudes
(and so position along a coastline) and depths may suggest
where to place fishing restrictions, whereas understanding
how such patterns vary across time may suggest when to
restrict fishing. These easy-to-collect variables are also likely
to be the most relevant to spatio-temporal management, for
example, one is unlikely to implement fishing restrictions
based on water temperature. Limiting the number of pre-
dictor variables also reduced collinearity between variables
(there were no interactions among the predictor variables)
and the possibility of model over-parameterization. We
Animal Conservation •• (2013) ••–•• © 2013 The Zoological Society of London
Mitigating small fish by-catch based on distribution patterns
measured depth as mean depth of a tow in meters (m).
Latitude (degrees north) was that of the vessel’s position at
the beginning of the tow. We measured time as a continuous
variable ‘day’, where 0 = first sample day = 25 September
2006. Since sampling started and ended with the fishing
season, progression of sampling days equates to progression
of the fishing season.
Analyses
We conducted analyses to determine the effects of depth,
latitude and time (and their interactions) on the dependant
variables (occurrence: presence/absence, density, LT).
Spatial and temporal autocorrelation are almost certain to
exist in data obtained from sampling that takes advantage
of fishing practices, as we have little control over when and
where sampling occurs. It was assumed that stations were a
random sample from the population. Thus, spatial and tem-
poral patterns were explored by analysing data with mixed-
effects models. The results of mixed-effects models can be
generalized to the entire population, have increased statisti-
cal power as a result of using all available data, and give a
flexible framework for analysing data with different types of
correlations (Ives & Zhu, 2006).
We conducted mixed-effects analyses using software pro-
vided in the packages lmer (binary variables) and lme
(continuous variables) within the statistical program R (R
Development Core Team, 2004). Fixed effects were latitude,
depth, day and their interactions. Station was treated as a
random effect to control for the non-independence of
repeated sampling of the same station. We verified all of the
models’ assumptions for ‘continuous response variables’,
that the residuals be normal, independent and identically
distributed, as per Pinheiro & Bates (2002). Non-normally
distributed data were transformed: log(square root) for
density and log for LT. We checked autocorrelation of
model residuals using the autocorrelation function and by
plotting a variogram for each of the dependent variables
(Pinheiro & Bates, 2002). Where autocorrelation of the
residuals was found, we modelled it with a simple moving
average model. We fit the model of presence/absence by
Laplace approximation, and the models of density and LT
using maximum likelihood. We selected the final model
using a sequential procedure for model selection, whereby
all covariates were included at first, and then possible
models of fixed effects were compared using likelihood-ratio
tests and model fit criteria (Hedeker 2005). We evaluated
candidate models using Akaike’s information criterion
(AIC). In the results for mixed-effects models (below), the
errors for covariates are standard errors. We present the
AIC value for the ‘best’ model (that with lowest AIC), as
well as the next smallest AIC value for comparison.
We also ran mixed-effects models using standardized pre-
dictor variables in order to compare the relative effects of
latitude, depth and day (each with different units of meas-
urement) in predicting occurrence, density and mean body
size of each taxon. The three predictor variables were
3
Mitigating small fish by-catch based on distribution patterns
rescaled by subtracting their respective means and dividing
by their standard deviations, thus converted to standard
deviation units.
Results
Diplectrum spp.
Accounting for the random effect of station in a mixed-
effects model revealed that occurrence of the Diplectrum
species increased primarily with increasing depth (esti-
mate = 0.20 ± 0.03, z = 5.83, P < 0.001) (AIC = 150.9 vs.
163.6). Occurrence also increased significantly with increas-
ing latitude (estimate = 2.31 ± 0.63, z = 3.68, P < 0.001), and
with the advancement of the fishing season (day: esti-
mate = 0.020 ± 0.004, z = 4.70, P < 0.001).
Mixed modelling also revealed that all three variables were
significant determinants of density for Diplectrum spp.
(AIC = −17.68 vs. −13.22). As with occurrence, depth was the
strongest predictor of density for this taxon (estimate =
0.010 ± 0.002, t = 5.93, P < 0.001, d.f. = 90), but latitude and
day also mattered although to lesser extents (latitude: esti-
mate = 0.11 ± 0.04, t = 2.66, P = 0.009, d.f. = 90; day: esti-
mate = 1.11E-03 ± 3.33E-04, t = 3.35, P = 0.001, d.f. = 90).
The mixed-effects model, with log(mean LT) per tow as
the dependent variable, only retained depth as an important
determinant of size for Diplectrum spp. (AIC = −343.3 vs.
−341.92). Mean size decreased with increasing depth for this
taxon (estimate = −1.03E-03 ± 4.00E-04, t = −2.48, P = 0.01,
d.f. = 92).
Prionotus stephanophrys
Occurrence of Pr. stephanophrys increased with increasing
depth (estimate = 0.17 ± 0.02, z = 7.33, P < 0.001), and also
with the advancement of the fishing season, although the
increase was only marginally significant (day: estimate =
0.007 ± 0.004, z = 1.99, P = 0.05) (AIC = 148.9 vs. 150.4).
Depth was the only significant determinant of density
for Pr. stephanophrys. Density increased with increasing
depth (estimate = 0.007 ± 0.003, t = 2.31, P = 0.03, d.f. = 45)
(AIC = 41.0 vs. 43.3).
Day and depth were retained as important determinants of
size for Pr. stephanophrys (AIC = −250.4 vs. −248.7). Mean
size decreased primarily with progression of the fishing
season (day: estimate = −1.74E-04 ± 7.44E-05, t = −2.33,
P = 0.02, d.f. = 44), but also with increasing depth (esti-
mate = −0.001 ± 4.00E-04, t = −3.03, P = 0.004, d.f. = 44).
Pseudupeneus grandisquamis
Occurrence of Ps. grandisquamis increased with the advance-
ment of the fishing season (day: estimate = 0.05 ± 0.01,
z = 5.83, P < 0.001) and with increasing depth (esti-
mate = 0.17 ± 0.04, z = 4.51, P < 0.001) (AIC = 190.1 vs.
211.0). Although day was more important than depth in
determining occurrence, their interaction was also statisti-
4 Animal Conservation •• (2013) ••–•• © 2013 The Zoological Society of London
S. J. Foster and F. Arreguin-Sánchez
cally significant for this species (estimate = −1.20E-
03 ± 3.00E-04, z = −4.36, P < 0.001).
Mixed modelling also revealed that latitude and day were
significant determinants of density for Ps. grandisquamis
(AIC = 86.5 vs. 88.1). Density increased with increasing lati-
tude (estimate = 0.25 ± 0.07, t = 3.32, P = 0.001, d.f. = 106),
and also as the fishing season progressed although to a
lesser degree (day: estimate = 1.50E-03 ± 4.00E-04, t = 3.42,
P = 0.001, d.f. = 106).
Depth and day were important determinants of mean size
for Ps. grandisquamis. Size of this species decreased with
the progression of the fishing season (day: estimate =
−9.02E-04 ± 8.00E-05, t = −10.73, P < 0.001, d.f. = 106), and
increased with increasing depth (estimate = 3.43E-02
± 1.80E-02, t = 1.91, P = 0.05, d.f. = 106) (AIC = −417.05 vs.
−415.04). An interaction between depth and latitude was also
important for this species (estimate = −1.62E-03 ± 8.05E-04,
t = −2.01, P = 0.05, d.f. = 106).
Stellifer illecebrosus
Accounting for the random effect of station revealed that all
three predictor variables, latitude, depth and day, were
important determinants of the presence of S. illecebrosus
across the study area (AIC = 205.8 vs. 207.3). Occurrence
decreased primarily as the fishing season progressed (day:
estimate = −0.015 ± 0.003, z = −4.54, P < 0.001) but also
occurred less frequently as depth and latitude increased
(depth: estimate = −0.09 ± 0.02, z = −4.54, P < 0.001; lati-
tude: estimate = −1.48 ± 0.56, z = −2.67, P = 0.008).
Mixed models also revealed that depth and day were
important determinants of density for S. illecebrosus
(AIC = 87.7 vs. 89.3). Depth was the most important
covariate, with density decreasing with increasing depth
(estimate = −0.013 ± 0.003, t = −3.65, P < 0.001, d.f. = 89).
Density also decreased as the fishing season progressed
(day: estimate = −0.0012 ± 0.0004, t = −2.94, P = 0.004,
d.f. = 89). Standardized regression coefficients for depth and
day were −0.14 ± 0.04 and −0.08 ± 0.02, respectively.
Finally, mean size of S. illecebrosus across the study area
was best predicted by a quadratic relationship with depth
(depth = 0.010 ± 0.004, t = 2.53, P = 0.01, d.f. = 92; depth2
= −1.64E-04 ± 7.73E-05, t = −2.12, P = 0.04, d.f. = 92)
(AIC = −289.7 vs. −287.4). Day was not an important pre-
dictor of size in this model, suggesting that the size of
S. illecebrosus was stable over the sampling period.
Among taxa comparison
The case study taxa differed in their spatial and temporal
distributions (Table 1). Latitude was an important predictor
of presence for Diplectrum and S. illecebrosus, although in
opposite ways, but was not important for Ps. grandisquamis
or Pr. stephanophrys. Whereas S. illecebrosus preferred
shallow waters, the three other taxa preferred deeper waters.
Occurrence of S. illecebrosus decreased over time across the
study site, but increased for the other taxa. Latitude was
positively related to densities of Diplectrum spp. and
S. J. Foster and F. Arreguin-Sánchez Mitigating small fish by-catch based on distribution patterns

Table 1 The direction of change in occurrence, density and mean size with increasing depth, latitude and progression of the fishing season for
four small fish taxa caught by shrimp trawlers in the southern Gulf of California, México
Diplectrum spp. Pr. stephanophrys Ps. grandisquamis S. illecebrosus
Occurrence Depth ↑* (2.72 ± 0.05) ↑* (63.42 ± 8.66) ↑ (0.71 ± 0.30 ) ↓ (−35.05 ± 8.64)
Latitude ↑ (1.25 ± 0.34) ↔ ↔ ↓ (−17.74 ± 6.65)
Day ↑ (1.25 ± 0.27) ↑ (44.25 ± 22.19) ↑* (1.26 ± 0.23) ↓* (−95.59 ± 21.04)
Density Depth ↑* (0.14 ± 0.02) ↑* ↔ ↓* (−0.14 ± 0.04)
Latitude ↑ (0.06 ± 0.02) ↔ ↑* (0.13 ± 0.04) ↔
Day ↑ (0.06 ± 0.02) ↔ ↑ (0.08 ± 0.02) ↓ (−0.08 ± 0.02)
Size Depth ↓* ↓ (−0.55 ± 0.18) ↑ (13.08 ± 6.85) Quadratic*
Latitude ↔ ↔ ↔ ↔
Day ↔ ↓* (−1.14 ± 0.45) ↓* (−5.84 ± 0.54) ↔

↑, significant increase; ↓. significant decrease; ↔. no significant change. Asterisk (*) indicates the spatio-temporal variable with greatest
influence on dependent variable, as determined by the relative magnitude of the standardized regression coefficients, in parentheses.

Ps. grandisquamis, but did not predict densities of
Pr. stephanophrys or S. illecebrosus. Depth was the most
important determinant of density for both Pr. stephanophrys
and S. illecebrosus, although in opposing directions. Densi-
ties of Diplectrum spp. and Ps. grandisquamis increased
over time, densities of S. illecebrosus decreased and those of
Pr. stephanophrys remained the same. Finally, whereas mean
size of Ps. grandisquamis and Pr. stephanophrys decreased
over the sampling period, mean sizes of Diplectrum spp. and
S. illecebrosus were stable over time.
The relative importance of spatial and temporal variables
in determining occurrence, density and mean size also varied
both within and among taxa (Table 1). Day was a significant
predictor of occurrence for all case study taxa. However, in
two taxa time mattered more than space whereas in the
other two the opposite was true. Either depth or latitude (or
both) were always more important than time in predicting
densities. Finally, whereas time was the most important
predictor of mean size for two taxa, depth was the key
predictor for the others.
The one consistent trend was that all taxa showed some
population size structure across the study site, with size
changing as a function of depth in all our study taxa
(Table 1).
Discussion
The results of our study suggested that any shrimp trawl
closures attempting to mitigate potential fishing effects on
small fishes from their incidental capture would need to be
permanent and biophysically diverse – there is no ‘one size
fits all’ solution. This recommendation is based on divergent
distribution patterns in time and space for all taxa, the
relative importance of spatial versus temporal variables, and
a relationship between body size and depth for all taxa. Our
results also provided insight into the potential for impact
from trawling on these by-catch taxa.
Evidence for fisheries impact
Our results provided only limited insight into the potential
for impact from trawling on these by-catch taxa. Prionotus
stephanophrys occurred more frequently across the study
site as the fishing season progressed, and although mean
body size decreased (present study) almost all captured indi-
viduals were mature (Foster & Vincent, 2010). Taken
together, the observed spatio-temporal distribution and
reproductive patterns for this species in the southern
Gulf suggested that the majority of individuals recruit to
the trawl grounds once mature. If this is indeed the case,
the trawl fishery might pose a problem as it removes indi-
viduals before they can spawn, but at the same time some
protection may be gained as the fishery appeared to leave
the youngest individuals alone. Pseudupeneus grandisquamis
also occurred more frequently and density increased over
time, accompanied by a decrease in population mean body
size (present study). But in this case, the proportion of
immature individuals increased over time across the study
site (Foster & Vincent, 2010), suggesting juveniles of this
species may have recruited into the study area during the
sample period. Understanding potential for impact of the
fishery on either of these species would require an under-
standing of whether juvenile survival or fecundity is the
limiting factor in its population viability (Walters & Martell,
2004).
There was no evidence for recruitment of S. illecebrosus
to the study site, with mean size of this species remaining
stable across the fishing season, but the results do suggest
potential for adverse fisheries impact. Stellifer illecebrosus
occurrence and density decreased over time, patterns
that may result from fishing pressure. The fact that
S. illecebrosus preferred shallow waters may also suggest
that the trawl fishery has a greater impact on this species,
compared with the other taxa. Although the industrial trawl
fishery is closed from April to August, small-scale boats
trawl year round (García-Caudillo & Gómez-Palafóx,
2005). These smaller boats cannot fish as deep as the indus-
trial ones and are thus more likely to affect shallow water
species. This suggests year-round trawling pressure may be
exerted on S. illecebrosus, whereas the other taxa might be
released from such pressure during the closed season. Other
fisheries operate in deeper waters during the summer
months, but they do not involve bottom trawling. On the
other hand, the fact that S. illecebrosus and the other case

Animal Conservation •• (2013) ••–•• © 2013 The Zoological Society of London 5

Mitigating small fish by-catch based on distribution patterns
study species are some of the more commonly caught fishes
in this fishery may suggest they are resilient to the trawl
pressure, although it is also possible they were caught in
much greater numbers in the past (Foster & Vincent, 2010).
To fully understand the potential impact of fishery on these
taxa, one would need to overlay spatio-temporal distribu-
tions of fishing effort on their distribution data (Andrew &
Pepperell, 1992) or monitor trends across several fishing
seasons.
We excluded from our analyses a physical variable, tem-
perature, which may actually be influential in species spatial
distribution. The reason we excluded it was that tempera-
ture was unlikely to be useful for the implementation of
spatio-temporal management. Temperature was however
predictably correlated with each of latitude, depth and time.
Our study suggested that temperature may be the driving
factor behind observed temporal changes in occurrence and
density of Diplectrum spp. Both occurrence and density
increased as the fishing season progressed, but with no con-
current change in mean size. Considering that latitude and
depth were also significant factors in determining both
occurrence and density of this taxon, observed changes over
time may be related to temperature increases in the more
northerly latitudes and at deeper depths as sampling pro-
gressed. Stellifer illecebrosus may also be subject to critical
temperature thresholds – southerly latitudes did favour this
species, which was also found more frequently and in
greater densities in shallower waters.
Mitigating impact
Our findings demonstrated the difficulties of regulating
shrimp trawl fisheries for particular by-catch species of
concern, in that their life history and distribution commonly
varied in a way that precludes narrowly focused remedial
action. Even the four taxa we analysed – a tiny sample of the
many small species obtained as by-catch in Mexican shrimp
trawls – exhibited distribution patterns that were at odds
with one another. If each of these species were of equal
conservation concern, then it would be hard to identify
specific spatial or temporal management measures to
support them all.
Indeed our results suggested that any restrictions on
shrimp trawling aimed at reducing potential impacts on the
case study taxa would need to be distributed in space, cov-
ering a range of latitudes and depths impacted by the
fishery, and be invoked throughout the fishing season. This
recommendation is based on: (1) divergent distribution pat-
terns in space and time, such that limiting fishery closures to
specific depths, latitudes or periods of time would not serve
all case-study taxa; (2) the relative importance of spatial
versus temporal variables, such that depth or latitude and
not time were the drivers of high densities; (3) the presence
of significant relationships between body size and depth,
such that limiting the fishery closures to specific depths
would only serve to protect a subsets of a species population
– trawl closures would instead need to be spatially diverse to
protect even one of the case study taxa.
6 Animal Conservation •• (2013) ••–•• © 2013 The Zoological Society of London
S. J. Foster and F. Arreguin-Sánchez
Apparent shifts in the space that individuals occupy as
they grow (or get older) are common in marine organisms,
and need to be addressed in management measures (Field
et al., 2006). Gradual movement to greater depths as indi-
viduals grow in size is common for shelf and slope species,
and so we might expect larger animals of such species to be
found in deeper waters. Pseudupeneus grandisquamis con-
formed to this expectation, but the patterns we observed for
Diplectrum spp. and Pr. stephanophrys were opposite to
those we expected, with larger individuals in shallower
waters. This could be explained by larger, mature individu-
als returning to shallow waters to spawn. Mean size of
S. illecebrosus was significantly related to depth, but the best
fitting model was one where larger animals were found at
mid-depths, whereas small animals were found in both
shallow and deeper waters. Further research is needed to
understand the underlying processes that produce these
patterns.
There are several approaches for deciding where to site
protected areas (e.g. Ban, Picard & Vincent, 2009), the dis-
cussion of which is outside the scope of this paper. But our
study suggests that when deciding when and where to imple-
ment trawl closures with an aim to mitigate potential
impacts on four taxa, let alone the full array of incidentally
captured small fishes, they should probably be permanent
and together encompass the full range of depths and lati-
tudes affected by the fishery. Such a proposal in support of
economically unimportant small fishes might seem problem-
atic were it not that most bottom trawling is intensive and
unsustainable in its direct and indirect effects on large
by-catch species, marine habitats and even target species
(e.g. Kaiser & Groot, 2000; Gillett, 2008). Closures would
have obvious benefits for most taxa, especially where they
occur in a context of declining target resources and increas-
ing overheads (e.g. burdensome fuel costs; Gillett, 2008;
Sumaila et al., 2008), as in the Gulf of California shrimp
fishery. Gear closures in the southern Gulf have the added
benefit of support from the majority of industrial trawl
fishers who were consulted about potential solutions for the
problems facing their fishery (Foster & Vincent, 2009).
Acknowledgements
This is a contribution from Project Seahorse. This study was
made possible thanks to collaborations with Dr. José Trini-
dad Nieto Navarro and his colleagues from the Escuela
Nacional de Ingeniería Pesquera (ENIP) in San Blas,
Nayarit, México. S.J.F. was supported by the International
Development Research Centre (Canada), IODE, Natural
Sciences and Engineering Research Council of Canada and
The University of British Columbia grants and fellowships,
and the John G. Shedd Aquarium in Chicago and Guylian
Chocolates Belgium through our partnerships for marine
conservation. F.A.-S. was supported by programmes from
the IPN. Fieldwork was coordinated by ENIP, and
CICIMAR was supported by grants from CONACyT-
SAGARPA (12004), SEP-CONACyT (104974), SIP-IPN
(20131266), EDI and COFAA. Many thanks to the techni-
S. J. Foster and F. Arreguin-Sánchez
cians and laboratory assistants who aided in sample and
data collection: J.A. Cardenas, D. Cisneros, E. Gallegos, A.
Hernández, S.P. Padilla, J.E. Rincones, and E.S. Sandoval;
and to P. Arcese, I. Caldwell, T. Carruthers, T. Jeanniard du
Dot, S. Hinch, M. Kaiser, P. Molloy, J. Rist, A. Trites,
A.C.J. Vincent, M. Yasue and four anonymous reviewers
for help with analyses and comments on the paper. Last but
not the least, our utmost thanks to the fishers who gener-
ously shared their fishing vessels with S.J.F. for the entire
2006–2007 shrimp fishing season.
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