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Biological Conservation xxx (2009) xxx–xxx

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Biological Conservation
journal homepage: www.elsevier.com/locate/biocon

Small cetacean captures in Peruvian artisanal fisheries: High despite

protective legislation
Jeffrey C. Mangel a,b,*, Joanna Alfaro-Shigueto a,b, Koen Van Waerebeek c, Celia Cáceres a,
Stuart Bearhop b, Matthew J. Witt b, Brendan J. Godley b
a
Pro Delphinus, Octavio Bernal, 572-5, Lima 11, Peru
b
Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall, TR10 9EZ, United Kingdom
c
Peruvian Centre for Cetacean Research (CEPEC), Museo de Delfines, Pucusana, Lima 20, Peru

a r t i c l e i n f o a b s t r a c t

Article history: We detail the first direct, at sea monitoring of small cetacean interactions with Peruvian artisanal drift
Received 16 April 2009 gillnet and longline fisheries. A total of 253 small cetaceans were captured during 66 monitored fishing
Received in revised form 17 August 2009 trips (Gillnet: 46 trips; Longline: 20 trips) from the port of Salaverry, northern Peru (8o140 S, 78o590 W)
Accepted 21 September 2009
from March 2005 to July 2007. The most commonly captured species were common dolphins (Delphinus
Available online xxxx
spp.) (47%), dusky dolphins (Lagenorhynchus obscurus) (29%), common bottlenose dolphins (Tursiops
truncatus) (13%) and Burmeister’s porpoises (Phocoena spinipinnis) (6%). An estimated 95% of common
Keywords:
dolphin bycatch was of long-beaked common dolphins (Delphinus capensis). Overall bycatch per unit
Bycatch
Artisanal fisheries
effort for gillnet vessels (mean ± sd) was estimated to be 0.65 ± 0.41 animals.set1 (range 0.05–1.50)
Small cetaceans and overall catch (bycatch and harpoon) was 4.96 ± 3.33 animals.trip1 (range 0.33–13.33). Based upon
Gillnets total fishing effort for Salaverry we estimated the total annual average small cetacean bycatch by gillnet
Longlines vessels as 2412 animals.year1 (95% CI 1092–4303) for 2002–2007. This work indicates that, in at least
Catch per unit effort one Peruvian port, bycatch and harpooning of small cetaceans persist at high levels and on a regular basis,
particularly in driftnet vessels, despite the existence since the mid-1990s of national legislation banning
the capture of marine mammals and commerce in their products. It is concluded that the coast of Peru is
likely still one of the world’s principal areas for concern regarding high small cetacean bycatch and there
is clearly an urgent need to increase the geographic scope of observer effort to elucidate the full magni-
tude of this issue.
Ó 2009 Elsevier Ltd. All rights reserved.

1. Introduction Fisheries bycatch has been of growing concern in recent dec-

ades (Brothers, 1991; Northridge, 1991; Perrin et al., 1994). The by-
Small scale coastal, or artisanal, fisheries make up the vast catch of long-lived, late maturing, low fecundity species like
majority of global fishers, produce about half of global annual fish marine mammals, seabirds, and sea turtles have received particu-
catch and provide most of the fish for human consumption in the lar attention and it is now clear that fisheries interactions pose
developing world (Berkes et al., 2001). These fisheries are typically one of the greatest risks to the survival of many populations
highly dispersed and are particularly prevalent in developing na- (Spotila et al., 2000; Lewison et al., 2004; Read, 2008). While initial
tions where regulations to monitor or manage these fisheries are attention was often primarily focused on bycatch by large indus-
frequently underdeveloped, unenforced or non-existent (Berkes trial fleets like tuna purse-seines, and high seas driftnets (Hall
et al., 2001). Despite their size and importance, however, artisanal et al., 2000), efforts have been intensifying to estimate the rates
fisheries remain under-studied in comparison with large scale and evaluate the impacts of bycatch in artisanal fisheries (D’Agrosa
industrial fleets (Berkes et al., 2001; Lewison et al., 2004; Pauly, et al., 2000; Moreno et al., 2006; Peckham et al., 2007, 2008; Alfaro
2006; Soykan et al., 2008). Shigueto et al., 2008) and those working in coastal seas (D’Agrosa
et al., 2000; Slooten, 2007).
* Corresponding author. Address: Pro Delphinus, Octavio Bernal, 572-5, Lima 11, Due to their circumglobal and coastal distributions, small ceta-
Peru. Tel.: +51 1 241 3081; fax: +51 1 241 3081. ceans are subject to human exploitation both from bycatch and di-
E-mail addresses: [email protected] (J.C. Mangel), [email protected] rect take (Jefferson and Curry, 1994; Reeves et al., 2003; Read et al.,
(J. Alfaro-Shigueto), [email protected] (K. Van Waerebeek), celiamaria81@
2006; Clapham and Van Waerebeek, 2007; Read, 2008). National
yahoo.es (C. Cáceres), [email protected] (S. Bearhop), [email protected]
(M.J. Witt), [email protected] (B.J. Godley).
and international legal measures to ban the take of dolphins and

0006-3207/$ - see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2009.09.017

Please cite this article in press as: Mangel, J.C., et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Con-
serv. (2009), doi:10.1016/j.biocon.2009.09.017
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2 J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx

porpoises in fisheries are meant to act as a protective measure to operated were voluntary participants in the project. Observers
reduce declines of cetacean populations (Northridge and Hofman, did not take part in fishing activity. Observers worked in all
1999). However, cetacean bycatch remains a concern worldwide months of the year over a total period of 29 months, in order to ac-
(Reeves et al., 2003; Lewison et al., 2004; Read et al., 2006; Read, count for possible seasonal variation in magnitude and spatial pat-
2008). Moreover, artisanal fisheries may contribute significantly terns of effort.
to cetacean mortality (Read et al., 2006). Gillnet fisheries in partic-
ular have been cited as probably the most significant cause globally 2.2. At sea observers
of small cetacean mortality (Jefferson and Curry, 1994; Dawson
and Slooten, 2005; Read et al., 2006). All observers were biologists and were trained in relevant data
Independent onboard observer programs have been widely collection methods including marine mammal identification. Data
used as an effective means to quantify bycatch (e.g. Gales et al., were gathered on specific gear used (longline or gillnet), the timing
1998; Beerkircher et al., 2002; Carretta et al., 2004; Rogan and and position (using GPS) of each set and any bycatch occurring. All
Mackey, 2007), and have been specifically recommended in the observers were equipped with cameras and photographed unusual
case of small cetacean captures in Peru (Reeves et al., 2005). Up- or unidentifiable captures for later species identification. Common
dated data on numbers of cetaceans caught and the spatio-tempo- dolphins Delphinus spp. were not identified to species in the boats,
ral distribution of cetaceans and bycatch are essential in defining nor were Tursiops truncatus assigned to inshore/offshore morpho-
the scale of any problem and in designing appropriate national type, considering there was a degree of uncertainty about positive
and regional management strategies (Reeves et al., 2003, 2005). identification among observers.
Moreover, the IUCN Cetacean Specialist Group (CSG) and the IWC Photos of Delphinus spp. (n = 38) examined by the authors indi-
Scientific Committee have both listed the Peruvian dusky dolphin cated bycatch of 36 long-beaked common dolphins Delphinus cap-
and Burmeister’s porpoise as priorities for cetacean bycatch ensis (94.7%) and two short-beaked common dolphins Delphinus
reduction. delphis (5.2%). This composition estimate is used in our extrapola-
In Peru, previous research into small cetacean captures has fo- tion to the wider estimate of take (Table 4). The overwhelming pre-
cused on the monitoring of landings of carcasses and fishmarkets ponderance of D. capensis found here is broadly consistent with the
for the presence of small cetacean products (Read et al., 1988; more than 99% of Delphinus catches belonging to D. capensis in Peru
Van Waerebeek and Reyes, 1990, 1994; García-Godos, 1992; Van based on a sample of 1067 common dolphins taken in coastal fish-
Waerebeek, 1994; Van Waerebeek et al., 1997, 2002; Majluf eries in the period 1984–1993 (Van Waerebeek, 1994).
et al., 2002). Captures of small cetaceans were thought to have
peaked in the period 1990–1993 when estimates of total take by 2.3. Shore-based observers
artisanal and commercial fisheries ranged between 15,000 and
20,000 animals per annum (Van Waerebeek and Reyes, 1994), Shore-based observers were employed in Salaverry to monitor
making it one of the largest small cetacean takes in the world. Min- daily fishing activity from September 2001 to March 2008. Observ-
isterial decrees (1990 and 1994) reinforced by a national law in ers collected data on the total number of fishing trips departing
1996 (Anonymous, 1996), prohibit the intentional take, landing and returning per day and per vessel type, locations of fishing
and sale of small cetaceans in Peru (reviewed in Van Waerebeek activity and associated catch and bycatch. Data collection was
et al., 1994), but this legislation is not fully enforced and the cap- based upon daily interviews with fishermen and monitoring of
ture and trade of small cetaceans continues (e.g. Van Waerebeek dockside activity. Respondents were informed that the information
et al., 2002). The legislation did, however, have the effect of reduc- would be kept anonymous and be used strictly for research pur-
ing reported landings and pushing the continuing trade in small poses. Fishermen returning from fishing trips were queried regard-
cetaceans into the black market which was much more difficult ing vessel type, fishing effort, target catch, and incidents of bycatch
to monitor (Van Waerebeek et al., 1997, 2002). In addition it was of small cetaceans, sea turtles or seabirds. Resulting data therefore
expected that, unlike before, at least some fishermen would simply are a census of fishing effort by gear type over the study period.
discard cetacean bycatch offshore so as to avoid any problems with
landings of legal fish catches. As a result, other methods are re- 2.4. Data analysis
quired to quantify the continuing catch of small cetaceans. Here
we report on recent at sea observations of artisanal gillnet and All observer data were managed in a Microsoft Access database.
longline activities allowing the first direct effort-corrected esti- Bycatch per unit effort (CPUE) was calculated on a per trip and per
mates of bycatch for artisanal fisheries operating from an impor- set basis for both fishing gears. For gillnet vessels, CPUE was also
tant Peruvian port. presented per length (km) and per area (km2) of net set. Descrip-
tive statistics are presented as mean ± standard deviation (SD) or
with 95% confidence intervals (CI) unless specified otherwise. Sta-
2. Materials and methods tistical tests were performed using SPSS 15.0 and Genstat 10. For
temporal analyses of total bycatch we used General Linear Models
2.1. Onboard observer scheme (GLMs) with normal errors, where CPUE was the dependent vari-
able with season and year as factors. In this instance CPUE was cal-
From March 2005 to July 2007 observers monitored a total of 66 culated on a trip by trip basis by dividing the number of bycatch
artisanal fishing trips (480 sets; 439 fishing days) for small ceta- incidents by the number of sets made. When it came to analysis
cean bycatch. Artisanal fisheries are defined here, according to of the bycatch for individual species the date distributions de-
Peruvian fisheries regulations, as containing boats with a maxi- parted significantly from normality and there were significant dif-
mum of 32.6 m3 of storage capacity, less than 15 m of length, ferences in variances among groups. We therefore used raw count
and principally based on the use of manual work during fishing data as our dependent variable, with number of sets included as a
operations (Ley General de Pesca, 2001). Trips monitored were covariate (to account for variation in effort across seasons/years)
on gillnet and longline vessels originating from the port of Sala- along with season and year as factors. We also employed GLMs
verry (8°140 S, 78°590 W), an artisanal port in northern Peru and for these analyses but fitted them with Poisson errors and a log link
home to over 100 fishing vessels (Alfaro-Shigueto et al., unpub- function. Season was divided as follows: season 1 = November–
lished results). Skippers (N = 21) upon whose vessels observers January; season 2 = February–April; season 3 = May–July; season

Please cite this article in press as: Mangel, J.C., et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Con-
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J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx 3

4 = August–October. All spatial analyses and maps were prepared face and were typically set in the afternoon and retrieved the fol-
using ESRI ArcMap 9.1, MATLAB 7.6 and Hawth’s Tools (Beyer, lowing morning. The average number of sets.trip1 was 7.4 ± 2.4
2004). Bathymetry values were determined with Global Gridded (range: 2–11). Total net length per set averaged 1948 ± 512 m
Relief Data (ETOPO2v2) with 2 minute resolution (USDOC, 2006). (range: 1097–3072). The only observed bait used was small ceta-
Quartic Kernel and 50% and 75% probability contour analyses were cean blubber or meat.
performed using 2 km grid spacing and least squares cross valida-
tion derived optimized smoothing factors for longline and gillnet
sets (25 km) and a smoothing factor of 35 km for small cetacean 3.2. Longline characteristics
capture locations.
A total of 20 trips by artisanal longline vessels (138 sets;
167,670 hooks; 129 fishing days) were monitored (Table S1). Six-
2.5. Estimating bycatch rates and total bycatch teen of 20 trips (80%) targeted dorado (Coryphaena hippurus) with
the remaining four trips targeting sharks (mainly blue and short-
Gillnet bycatch data for the study were grouped by month in or- fin mako). Mainlines for all trips were set at the sea surface and
der to derive monthly stratified CPUE estimates. These rates were were made of multifilament nylon rope. While trip lengths were
calculated in terms of catch.trip1, catch.set1, and to facilitate similar, vessels targeting sharks typically had more and longer sets
comparison with other studies, catch.km of net length1, and and deployed fewer, more widely spaced hooks than vessels tar-
catch.km2 of net area1 were also calculated. Given the small sam- geting dorado. Branchlines were made of narrow diameter nylon
ple size we did not prepare similar monthly stratified catch esti- multifilament cord, with branchline length of vessels fishing for
mates of longline bycatch. sharks approximately double that of vessels fishing for dorado.
Based upon the catch rates derived in this study and the data on Leader material used was either nylon monofilament when target-
monthly Salaverry fishing effort from 2002 to 2007, we were able ing dorado or metal cable when targeting sharks. Jumbo flying
to estimate the number of small cetacean captures (overall and per squid (Dosidicus gigas) was used as bait for both sharks and dorado
species) for the gillnet fleet. To derive these values we applied the while small cetacean blubber and meat was also used as bait by
monthly CPUE rates calculated in this study to the estimated num- vessels targeting sharks.
ber of monthly gillnet sets for the years 2002–2007. Monthly num-
ber of sets was estimated by multiplying the known number of
trips per month by the average number of sets per trip as deter- 3.3. Summary of small cetacean interactions
mined by this study. Month specific CPUE calculations were used
to generate an estimate. Bycatch data for each individual month A total of 253 dolphins and porpoises were observed by on-
were pooled to derive monthly CPUE values. As bycatch data were board observers as captured during the study period (Table 1). By-
left skewed, the monthly bycatch estimates were calculated by catch in gillnets accounted for 91.3% of all interactions recorded
multiplying the CPUE of sets with bycatch by the total number of with another 6.3% (n = 16), 2.0% (n = 5) and 0.4% (n = 1) coming
sets multiplied by the proportion of sets in that month estimated from longline harpooning, gillnet harpooning and longline bycatch,
to have bycatch (as determined in this study). Monthly catch esti- respectively.
mates for each year were then summed to arrive at annual totals.

3.3.1. Gillnets
3. Results Eighty percent of gillnet trips (37 of 46 trips; 104 sets) experi-
enced small cetacean bycatch and the majority of captures were
3.1. Gillnet characteristics of two species (common dolphins 50.2%; dusky dolphins 27.7%).
Captures also included common bottlenose dolphins (13.0%;
This project monitored 46 trips (341 sets; 319 fishing days) by n = 30), Burmeister’s porpoises (6.9%; n = 16), Risso’s dolphins
artisanal drift gillnet vessels (Table S1). A detailed summary of trip (Grampus griseus) (0.4%; n = 1), and unidentified small cetaceans
and net characteristics is presented in Table S1. All monitored trips (1.7%; n = 4; Table 2). Mean CPUE of small cetaceans was
targeted sharks and rays (mainly smooth hammerheads (Sphyrna 0.65 ± 0.41 animals.set1 (range: 0.05–1.50) or 4.96 ± 3.33 ani-
zygaena), eagle rays (Myliobatis spp.), blue sharks (Prionace glauca), mals.trip1 (range: 0.33–13.33) (Table S2). In addition to bycatch,
short-fin mako sharks (Isurus oxyrinchus) and thresher sharks three common bottlenose dolphins and two common dolphins
(Alopias vulpinus)). Gillnets observed were made of multifilament were harpooned for bait on three gillnet fishing trips by three dif-
nylon cord of varying mesh sizes. Nets were set at the ocean sur- ferent vessels.

Table 1
Species composition, capture methods and use of small cetacean carcasses of all interactions (n[%]) with gillnet and longline vessels. Percentages are read across for capture
methods and uses while species composition of gillnet bycatch subtotal and grand total (2nd and last columns from left) are tallied by column. The fate category ‘‘unknown”
refers to animals for which final fate was not recorded.

Species Grand Longline Longline Gillnet Gillnet bycatch

total harpoon bycatch harpoon
For bait Discarded Released Sold Eaten Eaten Unknown Gillnet
dead alive (boat) (home) subtotal
Delphinus spp. 120 (47) 2 (1.7) 0 2 (1.7) 22 (18.3)a 58 (48.3) 1 (0.8) 10 (8.3) 0 5 (4.2) 20 (16.7) 116 (50.2)
L. obscurus 73 (29) 8 (11.0) 1 (1.4) 0 35 (47.9)b 15 (20.5) 1 (1.4) 4 (5.5) 1 (1.4) 1 (1.4) 7 (9.6) 64 (27.7)
T. truncatus 33 (13) 0 0 3 (9.1) 10 (30.3) 17 (51.5) 1 (3.0) 0 0 0 2 (6.1) 30 (13.0)
P. spinipinnis 16 (6) 0 0 0 1 (6.3) 1 (6.3) 0 0 2 (12.5) 3 (18.8) 9 (56.3) 16 (6.9)
Unidentified 10 (4) 6 (60.0) 0 0 0 0 0 0 0 0 4 (40.0) 4 (1.7)
G. griseus 1 (0.4) 0 0 0 0 1 (100) 0 0 0 0 0 1 (0.4)
Total 253 (100) 16 (6.3) 1 (0.4) 5 (2.0) 68 (26.9) 92 (36.4) 3 (1.2) 14 (5.5) 3 (1.2) 9 (3.6) 42 (16.6) 231 (100)
a
Four animals sold to longline vessel while at sea.
b
Two animals given to another gillnet vessel and two stored for use on a subsequent longline trip.

Please cite this article in press as: Mangel, J.C., et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Con-
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4 J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx

Table 2
Estimated annual bycatch of small cetaceans by gillnet vessels for the port of Salaverry for the years 2002–2007, mean (CI). Values are derived from annually pooled monthly
estimates of bycatch and known levels of monthly fishing effort for the port. Presented estimates are of total estimated small cetacean captures and of the four most commonly
captured species.

Year # Trips Estimated # sets Total estimated bycatch D. capensis L. obscurus T. truncatus P. spinipinnis
2002 411 3054 2002 (845–3776) 690 (431–1011) 812 (189–1979) 189 (112–266) 191 (94–769)
2003 620 4607 3212 (1356–6047) 1168 (709–1713) 1284 (279–3188) 311 (187–435) 263 (148–982)
2004 421 3128 2118 (945–3839) 825 (437–1334) 759 (155–1845) 213 (98–328) 183 (86–629)
2005 572 4250 2518 (1247–4323) 1186 (680–1892) 619 (173–1368) 303 (129–477) 237 (134–677)
2006 593 4406 2636 (1278–4505) 1158 (619–1931) 773 (216–1719) 285 (129–441) 228 (115–756)
2007 492 3656 1987 (881–3330) 814 (372–1418) 662 (156–1666) 255 (158–352) 129 (56–385)
Average 518 3850 2412 (1092–4303) 973 (541–1550) 818 (195–1961) 259 (136–383) 205 (105–699)

3.3.2. Longlines 3.6. Temporal distribution

Small cetacean bycatch was only observed on one (5%) longline
fishing trip by a vessel targeting sharks and using small cetacean Total bycatch trip1 varied seasonally (F3,42 = 4.4, p = 0.009) but
meat as bait. The bycatch was of a dusky dolphin, the branchline not annually, nor was there a significant interaction between sea-
having been entangled around its flukes/tail stock. In addition, son and year (Fig. S2a). Post hoc Scheffe tests indicate that total by-
however, on three of four longline trips targeting sharks (15% of to- catch in season 4 (August–October) was much higher than in other
tal observed trips), dolphins were harpooned for bait (Table 1). months (means ± SE: season 1 = 0.19 ± 1.9; season 2 = 0.61 ± 0.14;
While we did not prepare monthly stratified catch estimates, the season 3 = 0.77 ± 0.15; season 4 = 1.18 ± 0.2). In the case of com-
overall interaction rate for longline vessels targeting sharks was a mon dolphin bycatch, both season (Wald = 11.12, df = 3, p = 0.11)
relatively high 4.25 ± 3.86 animals.trip1 (range: 0–8, n = 20) due and year (Wald = 8.75, df = 2, p = 0.13) have significant effects, with
to the common practice of harpooning dolphins for bait. number of sets and the season  year interaction having no signif-
icant influence (Fig. S2b). In this instance, pairwise comparisons
indicate that season 1 has lower bycatch than other
3.4. Fates of captured cetaceans seasons (p < 0.05. Estimated marginal means (EMMs) ± SE: season

All harpooned animals, both by gillnet and longline vessels,

were used as bait (Table 1). Also, the one dusky dolphin bycaught
by a longline vessel, while captured alive, was killed and used as
bait. Twenty-nine percent of gillnet bycatch was used as bait,
including 54.7% of dusky dolphins. Ninety-seven percent of gillnet
entangled animals were recovered dead. Of these, the most fre-
quent fate of the carcass was for it to be discarded at sea (39.8%).
Half of all common dolphins (50.0%) and a similar proportion of
common bottlenose dolphins (56.7%) bycaught in gillnets were dis-
carded dead. Gillnet entangled animals were also used for bait on
subsequent sets during the trip, later sold in local markets, con-
sumed on the boat or at home, released alive, or were given or sold
to other gillnet or longline vessels for use as bait. Although consti-
tuting a small part of the total, there is a suggestion that Burmeis-
ter’s porpoises may be preferred for human consumption, with
71.5% of known fate animals either consumed by the boat crew
or brought to shore to be eaten at home.

3.5. Spatial distribution

The scarcity of reliable bathymetry data in coastal zones

(<200 m) makes detailed interpretation of the depths of captures
difficult since most captures were in less than 250 m depth (Crac-
knell, 1999; Malthus and Mumby, 2003). However, several general
patterns do emerge when examining fishing effort and small ceta-
cean capture locations. Gillnet sets were more coastal than long-
line sets (Fig. 1a and b) with gillnet trips occurring over the
continental shelf and longline trips occurring on the continental
slope or pelagic. All small cetacean interactions appear to take
place on the continental shelf or near the slope (Fig. 1c). All har-
pooning and longline bycatch events occurred within their respec-
tive 90% probability contours of set locations. There was a
statistically significant difference in perpendicular distance to
shore of captures among the four most commonly taken species
Fig. 1. Set locations by (a) gillnet vessels, (b) longline vessels, and (c) of all gillnet
(H = 42.9, df = 3, Kruskal–Wallis, p < 0.001), with captures of Bur- bycatch. Also presented in each pane are 50% and 75% probability contours of
meister’s porpoises significantly nearer to shore than other species, fishing sets and gillnet bycatch (250 m, 750 m, 2000 m and 3000 m isobaths are
occurring in a small area fronting Salaverry (Fig. S1a–d). indicated).

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J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx 5

tions (Read et al., 1988; Van Waerebeek and Reyes, 1990; Van
Waerebeek, 1994; Van Waerebeek et al., 1997, 2002). Take ob-
served here consisted of the same species assemblage documented
in previous market studies (Read et al., 1988; Van Waerebeek and
Reyes, 1990, 1994; Van Waerebeek et al., 1997, 2002).

4.1. The magnitude of the issue

Our results indicate that, for this site, bycatch in gillnets is the
main cause of mortality with CPUE higher than published accounts
from the California driftnet fleet off the United States Pacific coast
(Barlow and Cameron, 2003), the Spanish driftnet fleet in the wes-
tern Mediterranean (Silvani et al., 1999), and Ecuadoran artisanal
gillnets (Félix and Samaniego, 1994) and comparable to CPUE for
Fig. 2. Monthly average number of trips by gillnet and longline vessels for the years the large scale Moroccan driftnet fleet in the southwest Mediterra-
2002–2007 determined from daily dock-side monitoring of fishing activity.
nean (Tudela et al., 2005). CPUE in the Irish driftnet fleet in the
northeast Atlantic was higher than observed here (Rogan and
1 = 0.67 ± 0.28; season 2 = 1.94 ± 0.38; season 3 = 2.59 ± 0.48; sea- Mackey, 2007), however total annual estimated catch was about
son 4 = 1.66 ± 0.44) and that bycatch in 2007 was much lower than half that we have estimated for the port of Salaverry. As in other
in 2006 and 2005 (p < 0.05. EMMs ± SE: 2005 = 2.23 ± 0.43; studies both in Peru (Ilo; Alfaro-Shigueto, unpublished results),
2006 = 2.39 ± 0.35; 2007 = 0.67 ± 0.28). Only year had a significant and in the southern ocean (Kock et al., 2006), South Georgia (Ash-
effect on dusky dolphin bycatch (Wald = 46.1, df = 3, p < 0.001). In ford et al., 1996), and Hawaii (Forney and Kobayashi, 2007) ceta-
this instance the highly significant effect is driven by multiple dif- cean bycatch rates at the vessel level in longlines were
ferences among seasons. Dusky dolphin bycatch was significantly considerably lower than those of gillnet vessels. The overall inter-
lower in seasons 1 (EMM ± SE = 0.13 ± 0.12) and 2 (0.44 ± 0.17) action rate for longline vessels in Peru has the potential to be high,
than in seasons 3 (1.5 ± 0.33) and 4 (4.37 ± 0.74) (Fig. S2c). In con- however, given the frequency of harpooning observed in Salaverry
trast, the bycatch of bottlenose dolphins (Fig. S2d) shows no strong (three of four trips targeting sharks).
seasonal patterns but year (Wald = 7.5, df = 2, p = 0.025) and total The Peruvian artisanal fleet has more than doubled in size from
sets (Wald = 20.2, df = 1, p < 0.001) are both significant, with higher 1997–2005 to 9667 vessels and vessels in the port of Salaverry rep-
bycatch in 2007 (EMM ± SE = 0.97 ± 0.37) than in 2006 resent only ca. 1% of that fleet and ca. 2% of gillnetters (Escudero,
(0.32 ± 0.11) and 2005 (0.34 ± 0.14). For Burmeister’s porpoises 1997; Estrella et al., 1999, 2000; Estrella, 2007). It is feasible there-
(Fig. S2e) there were no bycatch incidents observed in 2006 and fore that, at the national level, interactions between artisanal fish-
2007 so year is not included in the analysis. The bycatch of this eries and small cetaceans remain globally significant. Indeed, it is
species shows a very weak seasonal effect (Wald = 6.2, df = 2, conceivable that total mortality by the artisanal fishery is of the or-
p = 0.047) with lower bycatch in seasons 2 (EMM ± SE = der or greater than that estimated in 1990–1993 (15,000–20,000
0.40 ± 0.28) and 3 (0.33 ± 0.33) than in seasons 1 (2.00 ± 0.70) small cetaceans annually for all of Peru, Van Waerebeek and Reyes,
and 4 (2.00 ± 0.70). However, the sample size for this last species 1994). An annual catch rate of this magnitude would be one of the
is small and thus these results should be treated with some highest estimated takes globally, on the order of that reported for
caution. Japan, Sri Lanka, or the large scale Morrocan driftnet fleet (Bjorge
et al., 1991; Leatherwood, 1994; Reeves et al., 2003; Tudela et al.,
2005). For the port of Salaverry alone, our estimate of small ceta-
3.7. Estimating annual totals
cean captures is approximately equivalent to that of all recorded
fisheries in the United States of America (Read et al., 2006).
Based upon daily shore-based monitoring of fishing effort in
Salaverry we determined that there were an average of
4.2. Challenges to and opportunities for take reductions
518.2 ± 90 gillnet trips (range: 411–620 trips.year1) and
300.7 ± 25.2 longline trips (range: 272–341 trips.year1) per an-
Almost all gillnet bycatch was recovered dead and approxi-
num, for the years 2002–2007 (Table 2, Fig. 2). For the years
mately 40% of all entangled small cetaceans were discarded at
2002–2007 the estimated annual number of small cetaceans by-
sea. Thus, while 60% of carcasses were used opportunistically as
caught by gillnet vessels in the port of Salaverry was 2412 (95%
bait or for consumption, the fact that the other 40% of all bycatch
CI 705–4415). The number of small cetaceans harpooned by gillnet
was discarded indicates that interactions with small cetaceans
vessels was estimated to be on the order of some tens of animals.
are often unwanted. This also points to the mixed success of Peru’s
protective legislation. That legislation succeeded in reducing land-
4. Discussion ings and shrinking the market for small cetacean products, but
does not appear to have reduced small cetacean captures at sea.
The work presented here provides the first direct, at sea moni- Current practice stands in sharp contrast with the 1985–1994 sit-
toring of small cetacean interactions with Peruvian artisanal gillnet uation when discards were rare and most carcasses were landed to
and longline vessels. It has shown that, in at least one port in be sold, openly or covertly (e.g. Van Waerebeek and Reyes, 1994).
northern Peru, a sizeable level of bycatch, direct take through har- This suggests that the promotion and implementation of bycatch
pooning, and consumption of small cetaceans, continue despite the avoidance measures in the gillnet fishery may now, perhaps for
existence since the mid-1990s of national legislation banning the the first time, be acceptable to fishermen as a means of reducing
capture of marine mammals and commerce in their products. Pre- unwanted catch. Given prevailing levels of poverty, the extent
vious work monitoring the take of small cetaceans in Peru’s arti- and size of the fishery and the resources available for natural re-
sanal fisheries focused largely on dock-side monitoring of source management, closure of fishing areas to gillnetting or mod-
landing, monitoring of fishmarkets for small cetacean products ification of gillnets (Dawson, 1991) appear unimplementable. The
and assessing beach cast carcasses for evidence of fishery interac- use of acoustic alarms has been shown to have potential in

Please cite this article in press as: Mangel, J.C., et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Con-
serv. (2009), doi:10.1016/j.biocon.2009.09.017
 ARTICLE IN PRESS

6 J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx

reducing gillnet bycatch in some cetacean populations (Kraus et al., While large, the artisanal fishery is one of several fisheries oper-
1997; Kastelein et al., 2001; Barlow and Cameron, 2003; Cox et al., ating in Peruvian waters and potentially interacting with small
2003; Koschinski et al., 2006; Leeney et al., 2007) and should be tri- cetaceans. One must also consider interactions with other fisheries,
aled in the Peruvian gillnet fishery. most notably industrial and artisanal purse-seine vessels targeting
Clearly though, a demand for small cetacean products in the small schooling fish, especially anchovy. Based upon onboard ob-
form of bait and meat persists. Bait was collected from entangled server effort of 2% of the industrial fleet in 2002, van Oordt and
animals but also from animals harpooned specifically to collect Alza (2006) reported an average capture rate of 0.041 dol-
bait. Harpooning for bait occurred on both gillnet and longline ves- phins.set1. They noted that small cetacean captures in the fishery
sels. When used in gillnets, pieces of dolphin blubber and meat could be significant given the estimated 80,000 fishing trips.year1.
were tied to the center of each net pane. Dolphin blubber and meat Data on fishing effort for all fisheries operating in Peru’s coastal
was the only bait observed used in gillnets during the study and waters need to be compiled in order to more effectively set the re-
was used specifically due to its claimed effectiveness in attracting search agenda towards building a clearer understanding of the pos-
blue and short-fin mako sharks. Use of small cetaceans as bait was sible impacts on small cetacean populations.
also reported during interviews with fishermen (both in Salaverry
and Pucusana) where they noted dolphin meat’s particular effec- Acknowledgements
tiveness for catching sharks given its high blood and fat content
and its characteristic, unlike some fish bait, to remain intact and at- We kindly acknowledge the fishermen and field observers for
tached even after extended periods of soaking (this study; Van their generous collaboration. Pro Delphinus staff biologists and
Waerebeek, unpublished results). Previous work also reported this veterinarians P. Melly, M. Pajuelo, and D. Vega provided superb
usage and warned that increasing demand for small cetacean meat data and field work. We acknowledge also the assistance of the
and blubber as shark bait could offset any reductions in small ceta- Faculty of Fisheries, Universidad de Trujillo. We also extend our
cean take as a result of the ban on capture and commerce (Van thanks to the editor and two anonymous reviewers who provided
Waerebeek et al., 1997, 2002). The use of small cetaceans as bait important comments and suggestions on the manuscript. Pro Del-
has also been reported in coastal communities inter alia in Colom- phinus (JCM, JAS, CCB) research on cetacean bycatch has received
bia (Mora-Pinto et al., 1995; Avila et al., 2008), Argentina (Goodall funding from the Oak Foundation through Duke University, Rufford
et al., 1994), Chile (Lescrauwaet and Gibbons, 1994), Mexico Foundation, Whale and Dolphin Conservation Society, Cetacean
(Zavala-González et al., 1994) and the Philippines (Dolar, 1994), Society International and Cleveland Metroparks Zoo. CEPEC
but the practice is common worldwide. In discussions with fisher- (KVW) research on cetacean bycatch has benefitted from
men during this study regarding their use of small cetaceans for long-term International Fund for Animal Welfare (IFAW) support.
bait, a large number indicated that one reason for the use of dol- JCM and JAS are Overseas Research and Students Awards
phins and porpoises was the high cost of their preferred traditional Scheme (ORSAS) and University of Exeter scholarship awardees,
bait fishes like mackerel (Scomber japonicus). Although challenging, respectively. SB, BJG and MJW receive funding from Darwin Initia-
finding an appropriate, low-cost substitute bait to cetacean meat tive, Natural Environment Research Council and the South West
and blubber may reduce harpooning. This is particularly urgent gi- Regional Development Agency through the Peninsular Research
ven recent evidence that the practice of harpooning small ceta- Institute for Marine Renewable Energy.
ceans for use as longline bait is prevalent along the entire
Peruvian coast, most recently being reported in the southern port
of Ilo in November 2008 (Bernedo, personal communication). Appendix A. Supplementary material

4.3. Future directions Supplementary data associated with this article can be found, in
the online version, at doi:10.1016/j.biocon.2009.09.017.
The current study makes clear that small cetacean bycatch and
direct take continues despite the existence of national legislation References
prohibiting capture and commerce in their products. Our results
Alfaro Shigueto, J., Mangel, J.C., Seminoff, J.A., Dutton, P.H., 2008. Demography of
mandate renewed interest on the part of all stakeholders to expand
loggerhead turtles Caretta caretta in the southeastern Pacific Ocean: fisheries-
the scope of research and monitoring of small cetacean popula- based observations and implications for management. Endangered Species
tions and their interactions with Peru’s artisanal fleet. Our study Research 5, 129–135.
demonstrates the feasibility and use of independent observer pro- Anonymous, 1996. Ley 26585. Declaran a delfines y otros mamíferos marinos como
especies legalmente protegidas. Diario El Peruano, Normas Legales, Lima, 9 de
grams onboard artisanal fishing boats, and we strongly recom- Abril de 1996, 138665.
mend that such surveys be continued and expanded throughout Ashford, J.R., Rubilar, P.S., Martin, A.R., 1996. Interactions between cetaceans and
all fisheries of concern across the full geographic scale. Priority longline fishery operations around South Georgia. Marine Mammal Science 12
(3), 452–457.
should be given to increased monitoring of gillnet fisheries in the Avila, I.C., Garcia, C., Bastidas, J.C., 2008. A note on the use of dolphins for bait in the
center and north of the country where the fleet is concentrated. Gi- artisanal fisheries off Bahía Solano, Chocó, Columbia. Journal of Cetacean
ven the large number of ports and landing sites used by the arti- Research and Management 10 (2), 179–182.
Barlow, J., Cameron, G.A., 2003. Field experiments show that acoustic pingers
sanal fleet it may be more practicable to choose a number of reduce marine mammal bycatch in the California drift gill net fishery. Marine
‘index’ ports distributed along the coast and to focus on maximiz- Mammal Science 19 (2), 265–283.
ing onboard observer coverage in these locations. Observer effort Beerkircher, L., Cortes, E., Shivji, M., 2002. Characteristics of shark bycatch observed
on pelagic longlines off the Southeastern United States, 1992–2000. Marine
should optimally be continuous in order to account for any tempo- Fisheries Review 64 (4), 40–49.
ral variations in interactions, or should at least ensure an adequate Berkes, F., Mahon, R., McConney, P., Pollnac, R., Pomeroy, R., 2001. Managing Small-
coverage of all seasons. Special attention should be paid to interac- Scale Fisheries: Alternative Directions and Methods. International Development
Research Centre, Ottawa. 309pp.
tions with dusky dolphins and Burmeister’s porpoises since previ-
Beyer, H.L., 2004. Hawth’s Analysis Tools for ArcGIS. <http://
ous research indicate that the Peruvian populations of these www.spatialecology.com/htools>.
species form reproductively and genetically isolated stocks that Bjorge, A., Brownell, R.L., Perrin, W.F., Donovan, G.P., 1991. Significant direct and
should be subject to stock specific management measures (Van incidental catches of small cetaceans. Report of the International Whaling
Commission 42, 178–255.
Waerebeek, 1992, 1993; Cassens et al., 2003, 2005; Rosa et al., Brothers, N., 1991. Albatross mortality and associated bait loss in the Japanese
2005). longline fishery in the Southern Ocean. Biological Conservation 55, 255–268.

Please cite this article in press as: Mangel, J.C., et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Con-
serv. (2009), doi:10.1016/j.biocon.2009.09.017
 ARTICLE IN PRESS

J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx 7

Carretta, J.V., Price, T., Petersen, D., Read, R., 2004. Estimates of marine mammal, sea Malthus, T.J., Mumby, P.J., 2003. Remote sensing of the coastal zone: an overview
turtle, and seabird mortality in the california drift gillnet fishery for swordfish and priorities for future research. International Journal of Remote Sensing 24
and thresher shark, 1996–2002. Marine Fisheries Review 66 (2), 21–25. (13), 2805–2815.
Cassens, I., Van Waerebeek, K., Best, B., Crespo, E.A., Reyes, J., Milinkovitch, M.C., Mora-Pinto, D.M., Muñoz-Hincapié, M.F., Mignucci-Giannoni, A.A., Acero-Pizarro, A.,
2003. The phylogeny of dusky dolphins (Lagenorhynchus obscurus): a critical 1995. Marine mammal mortality and strandings along the Pacific coast of
examination of network methods and rooting procedures. Molecular Ecology Columbia. Report of the International Whaling Commission 45, 427–429.
12, 1781–1792. Moreno, C.A., Arata, J.A., Rubilar, P., Hucke-Gaete, R., Robertson, G., 2006. Artisanal
Cassens, I., Van Waerebeek, K., Best, P.B., Tzika, A., van Helden, A., Crespo, E.A., longline fisheries in Southern Chile: lessons to be learned to avoid incidental
Milinkovitch, M.C., 2005. Evidence for male dispersal along the coasts but no seabird mortality. Biological Conservation 127 (1), 27–36.
migration in pelagic waters in dusky dolphins (Lagenorhynchus obscurus). Northridge, S. 1991. Driftnet Fisheries and their Impact on Non-target Species: A
Molecular Ecology 14, 107–121. Worldwide Review. FAO Fisheries Technical Paper no. 320. FAO, Rome. 115 pp.
Clapham, P., Van Waerebeek, K., 2007. Bushmeat and bycatch: the sum of the parts. Northridge, S.P., Hofman, R.J., 1999. Marine mammal interactions with fisheries. In:
Molecular Ecology 16, 2607–2609. Twiss, J.R., Jr., Reeves, R.R. (Eds.), Conservation and Management of Marine
Cox, T.M., Read, A.J., Swanner, D., Urian, K., Waples, D., 2003. Behavioural response Mammals. Smithsonian Institution Press, Washington, D.C., pp. 99–119
of bottlenose dolphins, Tursiops truncatus, to gillnets and acoustic alarms. (471pp.).
Biological Conservation 115, 203–212. Pauly, D., 2006. Major trends in small scale marine fisheries, with an emphasis on
Cracknell, A.P., 1999. Remote sensing techniques in estuaries and coastal zones – an developing countries and some implications for the social sciences. Maritime
update. International Journal of Remote Sensing 20 (3), 485–496. Studies 4, 7–22.
D’Agrosa, C., Lennert-Cody, C.E., Vidal, O., 2000. Vaquita bycatch in Mexico’s Peckham, S.H., Maldonado Diaz, D., Walli, A., Ruiz, G., Crowder, L.B., Nichols, W.J.,
artisanal gillnet fishery: driving a small population to extinction. Conservation 2007. Small-scale Fisheries Bycatch Jeopardizes Endangered Pacific Loggerhead
Biology 14 (4), 110–1119. Turtles. PLoS ONE 2(10): e1041. 10.1371/journal.pone.0001041.
Dolar, L.L., 1994. Incidental takes of small cetaceans in fisheries in Palwan, central Peckham, S.H., Maldonado Diaz, D., Koch, V., Mancini, A., Gaos, A., Tinker, M.T.,
Visayas and northern Mindanao in the Philippines. Report of the International Nichols, W.J., 2008. High mortality of loggerhead turtles due to bycatch, human
Whaling Commission (Special issue 15), 355–363. consumption and strandings at Baja California Sur, Mexico, 2003–2007.
Dawson, S.M., 1991. Modifying gillnets to reduce entanglement of cetaceans. Endangered Species Research (5), 171–183.
Marine Mammal Science 7 (3), 274–282. Perrin, W.F., Donovan, G.P., Barlow, J. (Eds.), 1994. Report of the International
Dawson, S.M., Slooten, E., 2005. Management of gillnet bycatch of cetaceans in New Whaling Commission (Special issue 15). Gillnets and Cetaceans. International
Zealand. Journal of Cetacean Research and Management 7 (1), 59–64. Whaling Commission, Cambridge, UK. 629pp.
Escudero, L., 1997. Encuesta estructural de la pesquería artisanal del litoral Peruano. Read, A.J., 2008. The looming crisis: interactions between marine mammals and
Informe Progresivo del Instituto del Mar del Perú 59 (Junio). 87pp. fisheries. Journal of Mammalogy 89 (3), 541–548.
Estrella, C., 2007. Resultados generals de la segunda encuesta estructural de la Read, A., Van Waerebeek, K., Reyes, J., Mckinnon, J.S., Lehman, L.C., 1988. The
pesquería artisanal en el litoral Peruano: II ENEPA 2004–2005. 8pp. exploitation of small cetaceans in coastal Peru. Biological Conservation 46, 53–
Estrella, C., Guevara-Carrasco, R., Avila, W., Palacios, J., Medina, A., 1999. Informe 70.
estadístico de los recursos hidrobiológicos de la pesca artisanal por especies, Read, A.J., Drinker, P., Northridge, S., 2006. Bycatch of marine mammals in US and
artes, meses y caletas durante el primer semestre de 1999. Informe del Instituto global fisheries. Conservation Biology 20 (1), 163–169.
del Mar del Perú 148, 1–214. Reeves, R.R., Smith, B.D., Crespo, E.A., Notarbartolo di Sciara (compilers), G., 2003.
Estrella, C., Guevara-Carrasco, R., Avila, W., Palacios, J., Medina, A., 2000. Statistic Dolphins, Whales and Porpoises: 2002–2010 Conservation Action Plan for the
report on hydrobiological resources of artisanal fishery, by species, arts, ports, World’s Cetaceans. IUCN/SSC Cetacean Specialist Group, IUCN, Gland,
coves and months during the second semester 1999. Informe del Instituto del Switzerland and Cambridge, UK (ix+ 139pp.).
Mar del Perú 151, 1–194. Reeves, R.R., Berggren, P., Crespo, E.A., Gales, N., Northridge, S.P., Notarbartolo di
Félix, F., Samaniego, J., 1994. Incidental catches of small cetaceans in the artisanal Sciara, G., Perrin, W.F., Read, A.J., Rogan, E., Smith, B.D., Van Waerebeek, K., 2005.
fisheries of Ecuador. Report of the International Whaling Commission (Special Global Priorities for Reduction of Cetacean Bycatch. Report to the World Wide
issue 15), 475–480. Fund for Nature. 29pp.
Forney, K.A., Kobayashi, D.R., 2007. Updated estimates of mortality and injury of Rogan, E., Mackey, M., 2007. Megafauna bycatch in drift nets for albacore tuna
cetaceans in the Hawaii-based longline fishery, 1994–2005. NOAA Technical (Thunnus alalunga) in the NE Atlantic. Fisheries Research 86, 6–14.
Memorandum NMFS. NOAA-TM-NMFS-SWFSC-412. November. 35pp. Rosa, S., Milinkovitch, M.C., Van Waerebeek, K., Berck, J., Oporto, J., Alfaro-Shigueto,
Gales, R., Brothers, H., Reid, T., 1998. Seabird mortality in the Japanese tuna longline J., Van Bressem, M.F., Goodall, N., Cassens, I., 2005. Population structure of
fishery around Australia, 1988–1995. Biological Conservation 86, 37–56. nuclear and mitochondrial DNA variation among South American Burmeister’s
García-Godos, I., 1992. Captura estacional de cetáceos menores en la caleta de porpoises (Phocoena spinipinnis). Conservation Genetics 6, 431–443.
Ancón. Memoria X Congreso Nacional de Biología 02-07 agosto 1992, Lima, pp. Silvani, L., Gazo, M., Aguilar, A., 1999. Spanish driftnet fishing and incidental catches
273–279. in the western Mediterannean. Biological Conservation 90, 79–85.
Goodall, R.N.P., Schiavini, A.C.M., Fermani, C., 1994. Net fisheries and net mortality Slooten, E., 2007. Conservation management in the face of uncertainty:
of small cetaceans off Tierra del Fuego, Argentina. Report of the International effectiveness of four options for managing Hector’s dolphin bycatch.
Whaling Commission (Special issue 15), 295–304. Endangered Species Research 3, 169–179.
Hall, M.A., Alverson, D.L., Metuzals, K.I., 2000. By-catch: problems and solutions. Soykan, C.U., Moore, J.E., Žydelis, R., Crowder, L.B., Safina, C., Lewison, R.L., 2008.
Marine Pollution Bulletin 41 (1–6), 204–219. Why study bycatch? An introduction to the theme section on fisheries bycatch.
Jefferson, T.A., Curry, B.E., 1994. A global review of porpoise (Cetacea: Phocoenidae) Endangered Species Research 5, 91–102.
mortality in gillnets. Biological Conservation 67, 167–183. Spotila, J., Reina, R.D., Steyermark, A.C., Plotkin, P.T., Paladino, F.V., 2000. Pacific
Kastelein, R.A., de Haan, D., Vaughan, N., Staal, C., Schooneman, N.M., 2001. The leatherback turtles face extinction. Nature 405, 529–530.
influence of three acoustic alarms on the behavior of harbor porpoises Tudela, S., Kai Kai, A., Maynou, F., El Andalossi, M., Guglielmi, P., 2005. Driftnet
(Phocoena phocoena) in a floating pen. Marine Environmental Research 52, fishing and biodiversity conservation: the case study of the large-scale
351–371. Moroccan driftnet fleet operating in the Alboran Sea (SW Mediterranean).
Kock, K., Purves, M.G., Duhamel, G., 2006. Interactions between cetacean and Biological Conservation 121, 65–78.
fisheries in the Southern Ocean. Polar Biology 29, 379–388. USDOC, 2006. US Department of Commerce, National Oceanic and Atmospheric
Koschinski, S., Culik, B.M., Trippel, E.A., Ginzkey, L., 2006. Behavioral reactions of Administration, National Geophysical Data Center. 2-minute Gridded Global
free ranging harbor porpoises Phocoena phocoena encountering standard nylon Relief Data (ETOPO2v2). <http://www.ngdc.noaa.gov/mgg/fliers/06mgg01.
and BaSO4 mesh gillnets and warning sound. Marine Ecology Progress Series html>.
313, 285–294. van Oordt, F., Alza, L., 2006. Small cetacean interactions with the purse seine
Kraus, S.D., Read, A.J., Solow, A., Baldwin, K., Spradlin, T., Anderson, E., Williamson, J., fisheries off the Peruvian coast during 2002. Poster HCS201 In: Presented to the
1997. Acoustic alarms reduce porpoise mortality. Nature 388, 525. International Conference on the Humboldt Current System: Climate, Ocean
Leatherwood, S., 1994. Re-estimation of incidental cetacean catches in Sri Lanka. Dynamics, Ecosystem Processes and Fisheries. Lima, Peru, 27 November-1
Report of the International Whaling Commission (Special issue 15), 64–65. December (unpublished).
Leeney, R.H., Berrow, S., McGrath, D., O’Brien, J., Cosgrove, R., Godley, B., 2007. Van Waerebeek, K. 1992. Population Identity and General Biology of the Dusky
Effects of pingers on the behavior of bottlenose dolphins. Journal of the Marine Dolphin, Lagenorhynchus obscurus (Gray 1828) in the Southeast Pacific. Ph.D.
Biological Association of the United Kingdom 87, 129–133. Thesis, Institute for Taxonomic Zoology, University of Amsterdam. 160pp.
Lescrauwaet, A., Gibbons, J., 1994. Mortality of small cetaceans and the crab bait Van Waerebeek, K., 1993. Geographic variation and sexual dimorphism in the skull
fishery in the Magallanes area of Chile since 1980. Report of the International of the dusky dolphin Lagenorhynchus obscurus (Gray 1828). Fishery Bulletin 91,
Whaling Commission (Special issue 15), 485–494. 754–774.
Lewison, R.L., Crowder, L., Read, A.J., Freeman, S., 2004. Understanding impacts of Van Waerebeek, K., 1994. A note on the status of the dusky dolphins
fisheries bycatch on marine megafauna. Trends in Ecology and Evolution 19 (Lagenorhynchus obscurus) off Peru. Report of the International Whaling
(11), 598–604. Commission (Special issue 15), 525–527.
Ley General de Pesca, 2001. Reglamento de la ley general de pesca. Decreto Van Waerebeek, K., Reyes, J., 1990. Catch of small cetaceans at Pucusana port,
Supremo #012-2001-PE. Titulo 3, Capitulo 2, Articulo 30. central Peru, during 1987. Biological Conservation 51, 15–22.
Majluf, P., Babcock, E.A., Riveros, J.C., Arias Schreiber, M., Alderete, W., 2002. Catch Van Waerebeek, K., Reyes, J., 1994. Post-ban small cetacean takes off Peru: a review.
and bycatch of sea birds and marine mammals in the small-scale fishery of Report of the International Whaling Commission (Special issue 15), 503–
Punta San Juan, Peru. Conservation Biology 16 (5), 1333–1343. 519.

Please cite this article in press as: Mangel, J.C., et al. Small cetacean captures in Peruvian artisanal fisheries: High despite protective legislation. Biol. Con-
serv. (2009), doi:10.1016/j.biocon.2009.09.017
 ARTICLE IN PRESS

8 J.C. Mangel et al. / Biological Conservation xxx (2009) xxx–xxx

Van Waerebeek, K., Van Bressem, M.F., Reyes, J.C., Alfaro, J.A., Bello, R., Echegaray, Van Waerebeek, K., Alfaro-Shigueto, J., Montes, D., Onton, K., Santillan, L., Van
M., García-Godos, A., Ontón, K., 1994. Illegal Exploitation of Small Cetaceans in Bressem, M.F., Vega, D., 2002. Fisheries related mortality of small cetaceans in
Peru. Final Report to United Nations Environment Programme, Nairobi, and the neritic waters of Peru in 1999–2001. Document SC/54/SM10 Presented to
Whale and Dolphin Conservation Society, UK (76pp.). International Whaling Commission, Shimonoseki, Japan, May. 5pp.
Van Waerebeek, K., Van Bressem, M.F., Félix, F., Alfaro-Shigueto, J., García-Godos, A., Zavala-González, A., Urbán-Ramirez, J., Esquivel-Macías, C., 1994. A note on
Chávez-Lisambart, L., Ontón, K., Montes, D., Bello, R., 1997. Mortality of dolphins artisanal fisheries interactions with small cetaceans in Mexico. Report of the
and porpoises in coastal fisheries off Peru and southern Ecuador in 1994. International Whaling Commission (Special issue 15), 235–237.
Biological Conservation 81, 43–49.

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serv. (2009), doi:10.1016/j.biocon.2009.09.017