Journal of
Applied Ichthyology
J. Appl. Ichthyol. (2013), 1–7
© 2013 Blackwell Verlag GmbH
ISSN 0175–8659
Changes in fish bycatch during the shrimp fishing season along the eastern coast
of the mouth of the Gulf of California
By J. T. Nieto-Navarro1, M. Zetina-Rejón2, F. Arreguı́n-Sánchez2, D. S. Palacios-Salgado1 and F. Jordán3
1
Escuela Nacional de Ingenierı´a Pesquera, Universidad Autónoma de Nayarit, Bahı´a de Matanche´n, San Blas Nayarit, Me´xico;
2
Centro Interdisciplinario de Ciencias Marinas, Instituto Polite´cnico Nacional, La Paz, Baja California Sur, Me´xico; 3The
Microsoft Research – University of Trento Centre for Computational and Systems Biology, Rovereto, Trentino, Italy
Summary
One of the main concerns of shrimp fisheries is the associ-
ated impact on ecosystem biodiversity, particularly on fish
assemblages that are poorly characterized yet likely very rele-
vant to the health of the ecosystem. The continental shelf
along the eastern coast of the mouth of the Gulf of Califor-
nia is a region of high biodiversity that harbors highly pro-
ductive fisheries. This study aimed to analyze the changes in
the soft bottom fish assemblages caught as bycatch in the
shrimp fishery located in this region. Sampling was con-
ducted with commercial trawls at 16 fixed stations during the
2006–2007 shrimp-fishing season. A total of 103 fish species
from 80 genera and 47 families were collected. The ecological
and taxonomic diversity as well as the composition and
abundance of the fish community caught as bycatch in the
shrimp trawl fishery were found to be significantly different
in the autumn and winter. Ordination and similarity analyses
also revealed differing patterns. Species abundance was
strongly associated with sea bottom temperature, depth and
latitude. The effects of these abiotic variables on the
observed diversity patterns and the possible influences of the
fishery are discussed.
Introduction
The eastern coast of the mouth of the Gulf of California is
a highly productive area enriched by the influence of coastal
ecosystems, such as lagoons and estuaries (Ortiz-Pérez et al.,
2006), and by the confluence of the California Current (to
the south) and the North Equatorial Current (to the north;
Wyrtki, 1967; Kessler, 2006). This area supports several fish-
eries, most of which are artisanal. However, the most eco-
nomically relevant fishery in the region is the shrimp fishery
(Berdegué, 1976; Garcı́a-Caudillo and Gómez-Palafox,
2005). This is a sequential fishery in which artisanal fishers
exploit the inner waters, and the industrial trawl fleet
exploits waters of the continental shelf. One of the main
problems encountered by the shrimp fishery is the low selec-
tivity of trawl nets, which results in a high rate of fish and
invertebrate bycatch. The bycatch is commonly composed
of fish, crustaceans, mollusks, other marine animals and
dead matter. The worldwide average bycatch to shrimp ratio
is 20:1 (Eayrs, 2007), but this rate increases to 10 : 1 in the
shrimp fishery of the eastern coast of the Gulf of California
(Pérez-Mellado and Findley L. T, 1985). The catch composi-
tion has important implications for the sustainability of this
fishery, not only for the targeted shrimp stocks, but also for
associated commercial species and the ecosystem as a whole.
Received: May 15, 2011
Accepted: November 15, 2012
doi: 10.1111/jai.12160
For this reason, new approaches for management of fish
resources should consider the ecological effects of bycatch
(Kelleher, 2008). The impacts of trawling on benthic habi-
tats have been well documented and are related to the fre-
quency and severity of trawling. Effects of trawling on
benthic communities include reductions in organismal size,
diversity and biomass and usually require long-term recov-
ery times (Ball et al., 2000; Kaiser et al., 2002; Hiddink
et al., 2006).
Comparisons of fish community structures that highlight
the impacts of trawling using diversity and species richness
indices have received a great deal of attention, but no signifi-
cant differences in these indices have yet been detected
(Jennings and Reynolds, 2000). However, other approaches
have recently been developed to evaluate changes in diversity
using taxonomic diversity indices, which are more sensitive
to changes in perturbed habitats. The advantage of these
indices is that they are less dependent on size and sampling
effort and more sensitive to anthropogenic disturbances
(Warwick and Clarke, 1995; Leonard et al., 2006). Alterna-
tive non-parametric multivariate ordination analysis is
another useful tool for characterizing and discriminating
among sampling sites (Clarke and Warwick, 2001). The main
objectives of this study were to use ecological and taxonomic
descriptors to identify changes in the species composition of
fish caught as bycatch in the shrimp fishery and to analyze
how the patterns relate to abiotic conditions.
Materials and methods
Data
The study area was the continental shelf along the eastern
coast of the mouth of the Gulf of California. This area has
depths ranging from 10 to 59 m and is located on the shore-
line of the Southern Sinaloa and Northern Nayarit states
between 23.2°–22.2°N and 106.5° to 105.54°W (Fig. 1). The
area is characterized by soft sea bottoms that consist mainly
of sand and mud (Rodrı́guez-De la Cruz, 1981a,b).
Data were collected in four surveys carried out during the
2006–07 shrimp fishing season (September to March), two in
the autumn (September and November) and two in the win-
ter (January and February). The surveys were performed at
16 sampling stations distributed evenly throughout the study
area. Fish were collected in 64 trawls of 60 min duration at
a speed of 3 knots using commercial shrimp trawl nets (head
rope length: 28.96 m; mesh size in the cod end: 3.81 cm;
DOF, 1993). At each sampling station, surface and bottom
temperatures were recorded using a sensor HOBO Data
2 J. T. Nieto-Navarro et al.

(2006), considering six hierarchies (phylum, class, order, fam-

Mazatán
23.2° 16 ily, genus and species). We used weighted step lengths
15 between each taxonomic level, so that the distinctness of two
Mexico species connected at the highest taxonomic level was set to
13 Sinaloa 100. The estimated step length (16.7) was calculated by divid-
14
12
ing 100 by the number of hierarchies used in the analysis.
22.9° Therefore, wij = 16.7 for pairs of species in the same genus,
10
wij = 33.3 for species in the same family, etc. up to wij = 83.3
for species in the same class and wij = 100 for species in the
same phylum. The average taxonomic distinctness (Δ+) was

Te
8 9

aca
estimated using presence/absence data, and the number of

pan
22.6°
6 7 samples was not taken into account.
Pacific Ocean 5 4 Calculations of average taxonomic distinctness from 1000
simulated sub-samples of different numbers of species were
60 m

10 m
30 m

performed using the master list containing all fish species.

22.3°
3
11
1 distinctness values for all stations. This analysis addresses
Isobath
22°
106.5° 106°
Fig. 1. Study area along eastern coast of the mouth of Gulf of Cali-
fornia. Sampling stations (black dots) and isobaths are indicated
Logger (Onset Computer CorporationÓ 1995-2013, Bourne,
MA) with a data-storage feature equipped with an Optic
USB interface for data downloading (
0.2º accuracy).
In the laboratory, fish were identified to the species level
based on the taxonomic keys of Bussing and López (1993),
Fisher et al. (1995), and Robertson and Allen (2006) as well
as specific keys for the identification of the Diplectrum (Bor-
tone, 1977), Umbrina (Walker and Radford, 1992) and
Haemulopsis (López, 1981) genera.
Analysis
Numeric abundance was standardized to density using the
swept-area method (individuals per square kilometer; Sparre
and Venema, 1998), and distances were estimated using
a GPS. The fish community was analyzed in terms of Shan-
non’s diversity index (H’), Pielou’s evenness (J’) and average
taxonomic distinctness (Δ+; Clarke and Warwick, 1999) using
the equations:
X on the following variables: surface and bottom tempera-
H0 ¼  pi log2 pi
H0 H0
J0 ¼ ¼ (2)
Hmax InS
P P 
i\j wij
Dþ ¼  
SðS1Þ
 2 
where pi is the relative abundance of each species, calculated
as the proportion of individuals of a given species relative to
the total number of individuals in the community; Hmax is
the maximum possible value of the Shannon-Wiener diver-
sity, which is reached when the abundances of all species are
equal; S is the number of species; and wij is the weight given
to the path length linking species i and j in the hierarchical
classification. We used the classification proposed by Nelson
Nayarit
These distinctness values were used to produce 95% proba-
2 bility funnels, which were then plotted against the observed
Aguabrava lagoon
the question of whether Δ+ is different from the ‘expected
taxonomic distinctness’ based on a random selection from
the master list (Clarke and Warwick, 1998; Warwick and
105.5°
Clarke, 2001).
To test for significant differences between the autumn and
winter, we ran paired t-tests for ecological descriptors. A
non-metric multi-dimensional scaling (MDS) test was per-
formed using a Bray-Curtis similarity matrix estimated from
a log (x + 1) fish abundance matrix. MDS represents sam-
pling stations in 2-dimensional space such that the relative
distances between all points were in the same rank order.
Points that were close to each other represented sample sta-
tions that were similar in terms of taxonomic composition,
whereas points that were far apart represented different
assemblages (Clarke and Gorley, 2006).
Significant differences between multivariate groups were
assessed by a one-way analysis of similarities (ANOSIM;
Clarke, 1993). A similarity percentage (SIMPER) analysis
was used to identify those species responsible for the differ-
ences among groups (Clarke and Gorley, 2006). Species that
accounted for similarities greater than 90% were considered
key species. BIOENV, a method for correlating abiotic and
biotic data, was employed using Spearman’s rank correla-
tion coefficients (Clarke, 1993; Clarke and Warwick, 2001).
The biotic matrix was the same as that used in MDS,
ANOSIM and SIMPER, and the abiotic matrix was based
(1) tures, depth and the longitude and latitude of each sam-
pling station according to Munari and Mistri (2008). These
analyses were conducted using PRIMER 6.0 for Windows
software (PRIMER-E Ltd. Plymouth, UK).
Results
The temperatures at the bottom of the sea showed similar
patterns in the autumn and winter, such that higher tempera-
(3)
tures were generally recorded at sampling stations near to
the coast. The mean difference in temperature between the
autumn and winter was 5.61°C (Table 1).
In total, 16 542 individuals from 103 fish species were col-
lected; 7 of these species were Elasmobranchii, and 96 were
Actinopterygii. Species were grouped into 80 genera, 47 fami-
lies and 17 orders, of which Perciformes was the most abun-
dant, containing 51.5% of the species. Several species of the
following families were also observed: Carangidae, Scienidae,
Paralichthyidae, Haemulidae and Serranidae.
Fish by catch of the mouth of the Gulf of California 3

Table 1 4.4
Bottom (Sbt) and surface (Sst) sea temperatures recorded during
autumn and winter at each sampling station. Sample size (n) and 4.0
standard deviation (SD) are listed
3.6
Autumn Winter
3.2
Station Sbt n SD Sst* Sbt n SD Sst* H´
2.8
1 29.05 127 0.72 39.54 23.93 132 0.05 25.93
2 28.87 132 0.05 31.14 23.96 122 0.08 22.18 2.4
3 28.48 117 0.04 29.92 23.55 124 0.08 21.67
4 30.37 120 0.20 30.00 23.65 124 0.06 23.82 2.0
5 28.52 122 0.07 29.72 22.72 119 0.05 22.56
6 28.84 115 0.02 29.78 22.32 120 0.07 23.88 1.6
7 30.42 113 0.22 33.00 22.94 116 0.09 22.25
8 25.21 112 0.44 34.13 19.89 102 0.58 26.46 0.85
9 28.26 128 0.09 29.94 21.86 121 0.10 22.75
10 24.87 122 0.82 33.41 20.52 125 0.42 22.81
0.75
11 23.73 110 0.10 30.26 20.04 88 0.14 23.59
12 23.18 129 0.53 29.23 19.11 96 0.24 23.04
13 27.38 117 0.03 31.36 22.48 125 0.10 24.90 0.65
14 26.06 128 0.41 29.31 21.48 122 0.09 22.93
15 27.80 100 0.08 30.63 20.07 100 0.10 20.87 J´
16 27.34 122 0.14 31.16 20.12 121 0.30 22.75 0.55

*Corresponding to the first records of the HOBO data logger when

the net is at the sea surface. 0.45

Species richness values were higher in the winter (90) than
in the autumn (83); 70 species were present in both seasons.
The most numerically abundant species in the autumn were
Cathorops spp. (15.56%), Haemulopsis axillaris (13.73%),
Stellifer illecebrosus (8.85%) and Pomadasys panamensis
(7.1%). In the winter, Cathorops spp. (14%) was also among
the most abundant species, but other species were also very
abundant, including Pseudupeneus grandisquamis (11.4%),
Selene peruviana (10.9%) and Syacium ovale (10.4%).
The observed values of Shannon’s diversity index, evenness
and average taxonomic distinctness are shown in Fig. 2. Dif-
ferences in diversity were observed between the two seasons
analyzed; in the autumn, diversity values ranged from 2.1 to
4.1 bits/ind., whereas in the winter, diversity ranged from 1.8
to 3.8 bits/ind. Evenness values showed a similar pattern, with
a higher range of values observed in the autumn (0.45–0.82)
than in the winter (0.39–0.77). Taxonomic differences were
also observed between two seasons but in an inverse pattern to
that observed for diversity (Fig. 2). The t-test reveled signifi-
cant differences between the two seasons for the three ecologi-
cal attributes (H’, J’, Δ+: P = 0.005, P = 0.024 and P = 0.022,
respectively).
Funnel plots of average taxonomic distinctness are shown
in Fig. 3. Most stations fell within the 95% probability limits
of the simulation, which means that they did not depart sig-
nificantly from the null value. However, most observed val-
ues were below the expected mean, which indicates that the
fish community may be more taxonomically complex in the
winter than in the autumn, particularly near the sampling
stations that were relatively close to the shore (1, 2, 3, 13, 15
and 16).
The MDS ordination analysis detected two groups of sta-
tions from the data collected in the autumn (Fig. 4a); the
first group included sampling stations close to the coastline,
mostly in areas with depths up to 30 m (Group A: 1–7, 9
and 13), where 55.91% of the average similarity was defined
by 26 species [e.g. Cathorops spp. (7.21%), Stellifer illecebro-
sus (7.16%) and Larimus argenteus (6.27%)]. The second
group comprised sampling stations that were farther from
0.35
65
63
61
Δ+
59
Mean
57 Mean +-SE
55
Autumn Winter
Fig. 2. Box and whisker diagrams for Shannon’s diversity index
(H’), evenness (J’) and average taxonomic distinctness (Δ+) of the
soft bottom fish communities in autumn and winter
the coast, with depths mostly ranging from 30–60 min
(Group B: 8, 10–12, 14, 15 and 16). This group showed an
average similarity value of 58.36% and was mainly defined
by 23 species (e.g. P. panamensis (8.38%), Cyclopsetta querna
(8.37%) and S. ovale (6.83%)). The ANOSIM analysis indi-
cated significant differences between sampling stations from
Groups A and B (R = 0.729, P = 0.001). The SIMPER anal-
ysis identified 58 discriminant species with a mean dissimilar-
ity value of 54.56%. The main species contributing to the
dissimilarity were Eucinostomus currani (3.11%), Prionotus
stephanophrys (2.96%), H. axillaris (2.58%), S. ovale (2.5%)
and Centropomus robalito (2.47%).
The MDS analysis revealed three groups of sampling sta-
tions in the winter data (Fig. 4b). Group B remained con-
stant, with an average similarity value of 64.1% defined by
21 species (e.g. S. ovale (7.9%), P. grandisquamis (6.93) and
Diplectrum labarum (6.85%)). Group A remained nearly con-
stant, consisting of the same group of sampling stations with
the exception of 3 stations (1–3), which were placed into
Group C. The Group A sampling stations had an average
similarity value of 55.3% and were defined by 21 fish
4 J. T. Nieto-Navarro et al.

70 (a)
Phylum (1)
Class (1)
11 Order (8)
4
11 8
9 13 Family
7
16 10
9
6 8 5 (17)
12 6
14 14 15 Genus (27)
10 4
3 12 5
Δ + 60 2 1 7
1
13 Species (28)
3 15
2 16 (b)
Phylum (1)
Class (1)
Order (10)
50 Family (20)

20 30 40 Genus (23)

S Species (26)

Fig. 3. Funnel plot simulations of expected average taxonomic dis- Fig. 5. Comparison of taxonomic trees of soft bottom fishes col-
tinctness (bold line) for simulations of subsets of soft bottom fish lected at sampling station 16, during (a) autumn and (b) winter.
species. Limits within thin lines indicate 95% of simulated Δ+ values. Black dots = species remaining in the area during autumn and win-
Observed values for each sampling station are shown for autumn ter; grey dots = species entering the area in winter. Values in paren-
(black dots) and winter (white dots) theses = number of taxa

The ANOSIM analysis indicated significant differences

(a) Autumn among all paired comparisons for the different groups of
13 A sampling stations ([A,B] R = 0.90, P = 0.002; [A,C] R = 0.63,
3 B P = 0.012; and [B,C] R = 0.98, P = 0.008). The SIMPER
16
analysis identified 61 discriminant species for [A, B], with a
14
mean dissimilarity value of 57.4%. Species with the highest
10 15
percentage of dissimilarity were Cathorops spp. (3.33%),
9 1 2 P. stephanophrys (2.93%) and Synodus lacertinus (2.83%). In
12 5 contrast, for the [A, C] comparison, 53 discriminant species
8 were identified, with a mean dissimilarity of 51.3%. The sep-
7
11
aration of Groups A and C was mainly established by
4 Opisthopterus dovii (3.32%), Sphoeroides lobatus (3.13%) and
6 Bagre panamensis (2.83%). For [B, C] comparison, 49 discri-
minant species were identified, with a mean dissimilarity of
68.3%. Here, the most important species were S. illecebrosus
(3.67%), Synodus scituliceps (3.36%) and Cathorops spp.
(3.18%).
(b) Winter The results of the BIOENV analysis (Table 2) showed that
A
B in the autumn, the combination of variables having the
4 C
6 strongest correlation with fish species abundance was sea
bottom temperature (Sbt) in combination with depth (Dep)
7 and latitude (Lat). In the winter, the strongest correlation
8 5
was found with Sbt, followed by the combination of Sbt,
12
10 Dep. and Lat.
11
14 9
15 3
16 13 Discussion
1 2
The richness of fish species captured as bycatch in this study
(103) was below the lower range of that reported in previous
studies (105–187 species; Chávez and Arvizu J, 1972; Pérez-
Mellado and Findley L. T, 1985; van der Heiden et al., 1986;
Fig. 4. MDS ordination showing grouping of sampling stations Amezcua–Linares, 1987, 1996). This discrepancy can likely
during (a) autumn and (b) winter
be attributed to several different factors. First, the taxonomy
and nomenclature of certain genera observed in this study (e.
g. Sciades spp., Synodus spp. and Cathorops spp.) is still
species [e.g. P. grandisquamis (8.21%), Chloroscombrus orque- under revision and may have contributed to differences in
ta (7.73%) and Selene peruviana (6.88%)]. Sampling stations taxa aggregation (Kailola, 2004; Marceniuk, 2007; Mar-
in Group C had an average similarity of 60.6% and were ceniuk and Betancur, 2008). Additionally, the sampling sites
defined by 19 species [e.g. Anchoa walkeri (9.27%), S. illece- used in our study were closer to the coastline and located in
brosus (8.11%) and Symphurus elongatus (6.71%); Fig. 5]. shallower waters relative to those sampled by previous
Fish by catch of the mouth of the Gulf of California 5

Table 2 On the other hand, taxonomic diversity was inversely

Results of BIOENV analysis. Variables or combinations of variables related to ecological diversity, suggesting that community
that best correlate with the biotic matrix are shown in descending
structure became more heterogeneous due to a decrease in
order
ecological redundancy in the winter, as suggested by the
Season Variables rs higher values of taxonomic diversity observed. Although
some authors have argued that lower values of Δ+ are indica-
Autumn Sbt. Dep. Lat. 0.785 tive of more highly impacted areas (Warwick and Clarke,
Sbt. Dep. 0.739
Lat. 0.737 1998; Rogers et al., 1999), the increase in taxonomic diversity
Sbt. Dep. Lat. 0.721 observed in this study was a consequence of the species pres-
Sbt. Dep. Lat. Long. 0.712 ent due to the occurrence of cooler waters. Cooler waters are
Sbt. 0.705 correlated with the presence of species with wider
Sbt. Dep. Long. 0.697
distribution ranges and an affinity for lower temperatures,
Dep. 0.69
Sst. Sbt. Dep. Lat. 0.631 such as Antennarius avalonis, Hippoglossina tetrophthalma,
Sbt. Long. 0.624 Lophiodes caulinaris and Zalieutes elater. Additionally, the
Winter Sbt. 0.803 values of Δ+ were closer to the average expected mean found
Sbt. Dep. Lat. 0.8 during the winter, which suggests a less disturbed area that
Sbt. Lat. 0.775
Dep. Lat. 0.768 was influenced by the aforementioned fishing pattern. Similar
Sbt. Dep. 0.744 results were reported by Leonard et al. (2006), who noted
Sbt. Dep. Lat. Long. 0.735 that when the fishing effort of trawls was higher, values of Δ
Sst. Sbt. Dep. Lat. 0.719 + fell below the expected distribution and vice versa. In this
Sbt. Dep. Long. 0.691
case, sampling stations that fell below the expected distribu-
Sst. Sbt. Dep. Lat. Long. 0.679
Dep. Lat. Long. 0.671 tions are found in areas with a high trawling intensity due to
the high abundance of shrimp. In addition, sampling station
15 was close to the navigation channel and is expected to
suffer increased perturbation due to ship traffic and dredging.
authors. Furthermore, many of the previous studies were In these highly impacted areas, taxonomic homogenization
conducted two or more decades ago, and we do not account could eventually lead to an oversimplification of the commu-
for potential human or naturally induced perturbations that nity and similarity in terms of food resources (Olden et al.,
may have occurred since this time. 2004; Ramos-Miranda et al., 2005). This simplification may
The diversity patterns observed in this study were affect the structure (Jordán, 2009) and dynamics (Jordán
positively related to the sea bottom temperature. Amezcua– et al., 2008; Móréh et al., 2009) of the marine ecosystem.
Linares (1987)found a similar pattern of lower diversity in In the winter, taxonomic heterogeneity increased due to
the winter in an estuarine lagoon in this region. Based on the the presence of new species found only during this season.
BIOENV test, we also determined that latitude and depth These new species increased the taxonomic diversity by con-
are the most significant variables influencing fish community tributing a larger number of orders or families. To illustrate
structure. Munari and Mistri (2008) reported similar results this pattern, a comparison of the autumn and winter data
using the same test and determined that environmental and from sampling station 16 is shown in Fig. 5. The increase in
geographical variables influenced macrobenthic community taxonomic heterogeneity might also derive from the habitat
patterns. affinities of the different species (see FishBase: www.fish-
Although diversity patterns in the fish community were base.org). Group B remained constant between the autumn
clearly influenced by environmental conditions, abundance and winter, comprised of sampling stations far from the
and composition of bycatch were also related to fishing coastline with a depth range of 34–58 min and dominated
effort. When the shrimp abundance was higher at the by demersal as well as bentho-pelagic species. On the other
beginning of the shrimp fishing season, hauls were con- hand, the sampling stations closer to the coastline, with
ducted during the day, with a trawling duration of approxi- depths between 8.9 and 29.01 min, clustered into a single
mately 60–120 min. As the fishing season progressed and the group in the autumn (Group A) but fragmented into two
shrimp abundance decreased, trawling increased to approx. groups in the winter (Groups A and C) in consequence of
180–240 min (Pérez-Mellado and Findley L. T, 1985; Garcı́a- the disappearance of reef species, most likely due to second-
Caudillo and Gómez-Palafox, 2005). As a consequence of ary effects of the reduced temperature. However, some
the physical and biological impacts of trawling on the sea authors, including Wantiez (1992), suggest that this shift is
bottom, the abundance of shrimp and fish species decreased, associated with trophic migrations, the increase of pelagic
forcing fishers to trawl only at night. Crean and Symes species and the decrease of demersal fishes (Fig. 6). This
(1994) and Graham et al. (2007) noted that discard rates are model suggests that marine and pelagic species approach
correlated with fishing management strategies (or opera- the coast in the winter when perturbations occur at the
tions), which supports the idea that perturbations caused by beginning of the fishing season and leave adjacent habitat
increased fishing effort have a strong influence on the abun- available (Sainsbury, 1982; James et al., 1988; Wootton,
dance of bycatch. However, there are natural variations in 1990).
the abundance of demersal fish on soft bottoms that are We also observed changes in the abundance of several
modulated by environmental conditions, as reported by well-known pelagic species, such as Chloroscombrus orqueta
Mariscal-Romero and van der Heiden (2006) in the Central and Selene peruviana (Carangidae), which were the dominant
Mexican Pacific. These authors suggest that the changes in species in the winter but not in the autumn. Their abundance
oceanographic conditions generated by the California Cur- values increased from 2.09 to 9.20 and from 2.21 to 10.96,
rent, leading to cooler conditions in shallow waters, cause respectively, and contributed to the fragmentation of Group
some sensitive species to move into more protected areas. A into Groups A and C. These species use shoal and twilight
6 J. T. Nieto-Navarro et al.
100
80
Species (%)
60
40
20
0
AAutumn BAutumn AWinter BWinter CWinter
Sampling station group
Demersal Bentho-Pelagic
Pelagic Reef associated
Fig. 6. Ecological affinities of fish species according to their habitat
preference according to Fishbase (Froese and Pauly, 2008)
food habits (Helfman, 1978; Pitcher and Parrish, 1993).
These characteristics, coupled with the nocturnal activities of
the fishing fleet, contribute to the high abundance of these
species in trawl nets when a shoal is captured.
In general, the taxonomic, composition and abundance
patterns in the fish community caught as bycatch in the
shrimp trawl fishery depend on environmental conditions
such as temperature (González-Sansón et al., 1997) and the
location of the trawls (Chávez and Arvizu J, 1972; van der
Heiden, 1985). However, the disturbance caused by the
shrimp fishery and the exploitation strategies used by fishers
may regulate species abundance. We note that our study
reports certain taxonomic diversity patterns for the first time,
and for this reason, any comparison with previous studies on
diversity must be performed with care.
Acknowledgements
The authors would like to thank SEP-CONACyT (155900,
104974), SIP-IPN (20131458, 20131266), CONACyT reten-
tion (MOD-ORD-11-11-PCI-1103-03-12) and PROMEP
2011 for their institutional and financial support for the pro-
ject reintegration of former grantees 2011. The authors
would also like to thank EDI, COFAA and PIFI of the IPN
for financial support. We acknowledge the valuable com-
ments of the two anonymous referees who helped to improve
the manuscript.
Supporting Information
Additional Supporting Information may be found in the
online version of this article:
Table S1. Relative abundance (RA%) of soft bottom fishes
during autumn and winter, eastern coast of the mouth of the
Gulf of California. Resident species comprising 90% of the
similarity and discriminatory species for the conformation
and differentiation of the sampling station groups are shown.
Bold numbers = highest percentage of the species contribu-
tion to the similarity of each group. Dissimilarity: 6¼; Groups
of samples stations: A, B and C.
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Author’s address: J. T. Nieto-Navarro, Escuela Nacional de Inge-
nierı́a Pesquera, Universidad Autónoma de
Nayarit, Bahı́a de Matanchén km 12, Carretra a
los Cocos, San Blas Nayarit, México.
E-mail: nietojt@gmail.com