Tangled and drowned: A global review of penguin bycatch in fisheries

Abstract

Penguins are the most threatened group of seabirds after albatrosses. Despite being regularly captured in fishing gear, the threat to penguins, as a group, has not yet been assessed. We reviewed both published and grey literature to identify the fishing gear types that penguins are most frequently recorded in, the most impacted species and, for these susceptible species, the relative importance of bycatch compared to other threats. While quantitative estimates of overall bycatch levels are difficult to obtain, this review highlights that, of the world's 18 species of penguins, 14 have been recorded as bycatch in fishing gear and that gillnets, and to a lesser extent trawls, are the gear types that pose the greatest threats to penguins. Bycatch is currently of greatest concern for yellow-eyed Megadyptes antipodes (Endangered), Humboldt Spheniscus humboldti (Vulnerable) and Magellanic Spheniscus magellanicus penguins (Near Threatened). Penguins face many threats; reducing bycatch mortality in fishing gear will greatly enhance the resilience of penguin populations to threats from habitat loss and climate change that are more difficult to address in the short term. Additional data are required to quantify the true extent of penguin bycatch, particularly for the most susceptible species. In the meantime, it is crucially important to manage the fisheries operating within known penguin foraging areas to reduce the risks to this already threatened group of seabirds.

Full text

ENDANGERED SPECIES RESEARCH
Endang Species Res
Vol. 34: 373– 396, 2017
https://doi.org/10.3354/esr00869 Published November 30
INTRODUCTION
Penguins are one of the most threatened groups of
seabirds (Croxall et al. 2012), with 10 of the world’s
18 species listed as Vulnerable or Endangered on the
IUCN Red List and a further 3 deemed Near Threat-
ened (BirdLife International 2017a). Numerous ter-
restrial and marine threats contribute to this poor
conservation status. On land, erosion, grazing and
development resulting in habitat degradation, inva-
sive alien species, human disturbance and disease
are threatening several penguin species (García-Bor-
boroglu & Boersma 2013). At-sea threats include
habitat degradation due to industrial and commercial
© The authors 2017. Open Access under Creative Commons by
Attribution Licence. Use, distribution and reproduction are un -
restricted. Authors and original publication must be credited.
Publisher: Inter-Research · www.int-res.com
*Corresponding author: rory.crawford@rspb.org.uk
REVIEW
Tangled and drowned: a global review of penguin
bycatch in fisheries
Rory Crawford1,*, Ursula Ellenberg2,3, 4, Esteban Frere1,5, Christina Hagen6,
Karen Baird1, 7, Paul Brewin8, Sarah Crofts9, James Glass10, Thomas Mattern3, 4,
Joost Pompert11, Katherine Ross9, Jessica Kemper12, Katrin Ludynia13,14, Richard B.
Sherley15, Antje Steinfurth16, Cristián G. Suazo17,18, Pablo Yorio19, 20, Leandro
Tamini21, Jeffrey C. Mangel22, 23, Leandro Bugoni24, Gustavo Jiménez Uzcátegui25,
Alejandro Simeone26, Guillermo Luna-Jorquera27, Patricia Gandini5,
Eric J. Woehler28, Klemens Pütz29, Peter Dann30, Andre Chiaradia30, Cleo Small1
1BirdLife International Marine Programme, c/o RSPB, The Lodge, Potton Road, Sandy, Bedfordshire SG19 2DL, UK
Addresses for other authors are given in the Supplement at www.int-res.com/articles/suppl/n034p373_supp.pdf
ABSTRACT: Penguins are the most threatened group of seabirds after albatrosses. Although pen-
guins are regularly captured in fishing gear, the threat to penguins as a group has not yet been
assessed. We reviewed both published and grey literature to identify the fishing gear types that
penguins are most frequently recorded in, the most impacted species and, for these susceptible
species, the relative importance of bycatch compared to other threats. While quantitative esti-
mates of overall bycatch levels are difficult to obtain, this review highlights that, of the world’s 18
species of penguins, 14 have been recorded as bycatch in fishing gear and that gillnets, and to a
lesser extent trawls, are the gear types that pose the greatest threats to penguins. Bycatch is cur-
rently of greatest concern for yellow-eyed Megadyptes antipodes (Endangered), Humboldt
Spheniscus humboldti (Vulnerable) and Magellanic Spheniscus magellanicus penguins (Near
Threatened). Penguins face many threats; reducing bycatch mortality in fishing gear will greatly
enhance the resilience of penguin populations to threats from habitat loss and climate change that
are more difficult to address in the short term. Additional data are required to quantify the true
extent of penguin bycatch, particularly for the most susceptible species. In the meantime, it is cru-
cially important to manage the fisheries operating within known penguin foraging areas to reduce
the risks to this already threatened group of seabirds.
KEY WORDS: Fishery · Gillnet · Seabird · Trawl · Conservation · Direct mortality
O
PEN
PEN
A
CCESS
CCESS

Endang Species Res 34: 373– 396, 2017
development, as well as pollution and fisheries, both
indirectly through overfishing of prey species and
benthic habitat degradation from dredging or bottom
trawls, and directly through bycatch in fishing gear
(García-Borboroglu & Boersma 2013). The potential
impacts of climate change, both at sea and on land,
are threatening breeding habitats and prey availabil-
ity and are increasing the risk of disease (Trathan et
al. 2015).
The bycatch of penguins in fishing gear has been
the subject of few directed studies. Seabird bycatch
more broadly has been widely studied and identi-
fied as a substantial threat to affected species (Crox-
all et al. 2012). The interactions between Procellari-
iformes and longline fisheries, where birds are
accidentally captured on baited hooks (Brothers et al.
1999, Anderson et al. 2011), have been a focus of ap -
plied research. Furthermore, fatal interactions be -
tween large-winged seabirds (e.g. albatrosses) and
trawl fisheries have been identified (Weimerskirch
et al. 2000, Sullivan et al. 2006), due to the collision
of birds with net warp cables when foraging for dis-
cards behind vessels (Sullivan et al. 2006, Wat kins et
al. 2008) and net entanglement when trawl nets are
on the surface (Bull 2007, Pierre et al. 2013). More re -
cently, bycatch in purse seine fisheries has been
observed in Portugal (Oliveira et al. 2015) and Chile
(Suazo et al. 2016), principally affecting shearwa-
ters. Incidental capture in gillnet fisheries is esti-
mated to kill at least 400 000 seabirds annually, with
pursuit-diving seabirds identified as particularly
vulnerable (Žydelis et al. 2013).
Penguins, as pursuit divers, are most likely to inter-
act with gillnets (Žydelis et al. 2013) and trawls (Gan-
dini et al. 1999, González-Zevallos & Yorio 2006), al -
though they have also been recorded as bycatch in
longline fisheries in low numbers (Nel et al. 2002,
CCAMLR 2011). Penguin interactions with gillnets
are poorly understood, but entanglements are sus-
pected to occur when birds are actively foraging
(Simeone et al. 1999, Pütz et al. 2011), transiting or
resting on the surface (Majluf et al. 2002). Penguin
interactions with trawl fisheries have been primarily
recorded in Argentina, where Magellanic penguins
Spheniscus magellanicus are captured during the
haul as they attempt to feed on small non-target fish
that drop out of the net (González-Zevallos & Yorio
2006).
The limited published literature on penguin by -
catch is partially explained by the small-scale nature
of the fisheries with which penguins are most likely
to interact, particularly gillnet fisheries (Žyde lis et al.
2013). In general, such fisheries are highly dispersed
and diverse among jurisdictions, making monitoring
and enforcement difficult. For example, Peru, Chile
and Ecuador are together estimated to host around
90 000 artisanal fishing vessels, with boats fishing
from numerous locations along the Pacific coast of
South America (Goya et al. 2011).
In spite of the threatened status of many penguin
species (BirdLife International 2017a) and the clear
impact of bycatch on other seabirds (Anderson et al.
2011, Croxall et al. 2012), there has been no attempt
so far to collate the available data to understand the
potential impacts, knowledge gaps and conservation
priorities. Here, we aim to address this by reviewing
the published and grey literature to identify: (1) the
fishing gears of greatest concern with re gard to pen-
guin bycatch; (2) the species most at risk from bycatch
(and for these species, the relative importance of by -
catch compared to other threats); (3) the areas of con-
siderable penguin−fisheries interaction; (4) key con-
servation actions.
METHODS
A workshop on penguin bycatch at the 8th Inter-
national Penguin Conference in Bristol in 2013
brought together a network of experts that commit-
ted to collating all available knowledge on the topic.
The re sults of the extensive literature review on
global penguin bycatch were then discussed at a
follow-up bycatch workshop during the 9th Interna-
tional Penguin Conference in Cape Town in 2016.
The collated information includes published and
unpublished scientific papers and reports identified
via searches in Internet repositories (e.g. Google
Scholar), Web of Science and authors’ own (or asso-
ciated) unpublished works. The limited number of
publications on penguin bycatch (Figs. 1 & 2) means
that some older, potentially outdated, papers are re -
ferenced; thus some fisheries activities may have
changed in the intervening period. However, all
information presented here was included for its
broader relevance in understanding the scale and
species involved in penguin bycatch. Where bycatch
rates are available, these data and associated fleet-
wide bycatch estimates are included, although
many incidents are recorded simply as the number
of birds caught on a single trip.
This review is undoubtedly impacted by publishing
bias, and anecdotal reports of large bycatch events
are also more likely to be recorded than zero catches
(Wallace et al. 2010). In addition, studies are often not
conducted randomly, and may have been focussed in
374

Crawford et al.: Penguin bycatch in global fisheries
areas of perceived conflict or where resources hap-
pen to be available (Figs. 1 & 2). Irrespective, this re -
view will use the available information to broadly
identify the species, regions and gears of concern to
focus further efforts. Species’ ecology, fisheries active
in their range and consideration of the relative im -
portance of bycatch versus other threats are all fac-
tors used to determine future priorities.
Results are grouped into 4 regions: South America,
southern Africa, Australasia, and Antarctica includ-
ing the Southern Ocean islands. They are further
divided into countries where relevant. For each coun-
try, data are presented for the different fishing meth-
ods used. This approach allows cumulative impacts
for species across national jurisdictions to be consid-
ered based on all fishing methods in use, but also
results in the identification of relevant national-level
recommendations.
Penguin populations in the Antarctic and South-
ern Ocean islands were considered as a separate
region because of their low coastal fisheries effort,
as well as similar management regimes, particu-
larly under the Convention for the Conservation of
Ant arctic Marine Living Resources (CCAMLR). The
issues, species and management structures were
considered to be more relatable on this basis, rather
than attempting to link islands to distant continental
land masses.
375
Fig. 1. Penguin bycatch records in gillnets, trawls, longlines and purse seine fisheries. Records were assigned to the area from
which the data were obtained (where available; data without a specific location were assigned within the relevant species’
range) (coloured circles). The size of the circles indicates the relative number of records for that location. Source: Royal Society
for the Protection of Birds
0
5
10
15
20
25
30
35
40
Antarctica and
Southern Ocean
Islands including
CCAMLR area
Australasia
Southern Africa
South America
No. of publ. studies
(with different datasets)
No. of unpubl. studies/reports
No. of studies recording only
bycatch presence/absence
No. of studies recording only
number of birds bycaught
No. of studies providing by-
catch rate (birds/fishing effort)
No. of studies providing
bycatch estimate for fleet
Fig. 2. Summary of penguin bycatch records used in this re-
view (by region). Data are from published and unpublished
studies unless otherwise stated

Endang Species Res 34: 373– 396, 2017
RESULTS
South America
Breeding colonies of 7 species of penguins are
found in 4 countries (plus the Falkland Islands/Islas
Malvinas) in South America: Magellanic Spheniscus
magellanicus and southern rockhopper Eudyptes
chrysocome (both breed in Argentina, Chile and the
Falkland Islands/Islas Malvinas1), Humboldt S. hum-
boldti (Chile, Peru), Galápagos S. mendiculus (Ecua -
dor), macaroni E. chrysolophus and king Apteno dytes
patagonicus (both found in the Falkland Islands/Islas
Malvinas and Chile) and gentoo Pygoscelis papua
(Argentina and the Falkland Islands/Islas Malvinas)
(Kusch & Marin 2012, García-Borboroglu & Boersma
2013). During the non-breeding period (austral autumn
and winter), Magellanic penguins from colonies at
the Atlantic/Argentinean coast migrate through and
feed in the waters off Uruguay and Brazil.
Trawls and gillnets are the most important fishing
gears that penguins interact with in South America
(Table 1) (Simeone et al. 1999, González-Zevallos &
Yorio 2006, Žydelis et al. 2013). Due to the spatial
extent of these fisheries and differences in gear spec-
ifications and fishing operations, the interaction be -
tween fisheries and penguins — and the available in -
formation on these interactions — varies significantly
amongst the countries in South America.
Argentina
Trawl fisheries. Penguin bycatch in Argentina has
largely been recorded in trawl fisheries, particularly
focussed on Golfo San Jorge, Patagonia, which is
home to over 15% of the total Argentinean breeding
population of Magellanic penguins (Pozzi et al. 2015),
estimated at 950 000 birds (Schiavini et al. 2005).
The Argentine red shrimp Pleoticus muelleri double-
rig beam freezer trawl fishery active in Golfo San
Jorge was estimated, based on direct observation on
vessels, to kill 642 (±225) Magellanic penguins an -
nually in the late 1990s (Gandini et al. 1999). Further
observations, conducted between 2005 and 2007,
estimated 895 penguin mortalities fleet-wide across
the 3 years of the study period (González-Zevallos et
al. 2011). This study also highlighted that Magellanic
penguin bycatch was more likely closer to shore, and
that mortality increased with longer hauls (González-
Zevallos et al. 2011).
High seas ice trawlers target Argentine hake Mer-
luccius hubbsi in the same fishing area from Septem-
ber to May. Direct observation has given variable
estimates of fleet-wide Magellanic penguin mortality
over 2 yr: 35 deaths were estimated in 2003, and 1516
in 2004 (González-Zevallos & Yorio 2006). Although
there are no recent studies of this fishery, foraging
Magellanic penguins continue to overlap with shrimp
and hake vessels operating in adjacent waters during
the breeding season (Yorio et al. 2010).
Marinao & Yorio (2011) studied the incidental mor-
tality of seabirds in the coastal ice trawlers targeting
Argentine red shrimp in the Isla Escondida fishing
area, estimating 53 Magellanic penguins killed in
3149 hauls from 2006−2008, a mortality rate of 0.003
penguins haul−1. Penguin bycatch was also recorded
from 2008−2012 in both the coastal red shrimp and
hake fishery operating in the same area (Marinao et
al. 2014). In total, 203 adult Magellanic penguins were
incidentally caught in nets, occurring almost exclu-
sively in the hake fishery (97.5% of captures) and
largely occurring closer to penguin colonies (85% of
captures within 45 km of the coast; Marinao et al.
2014). Mean monthly capture rate was 0.087 birds
haul−1 (range 0.01−2.07 birds haul−1).
Lower levels of Magellanic penguin bycatch have
been recorded in the cornalito Odontesthes incisa
pelagic pair trawl fishery in Puerto Quequén (38° S,
58° W), with an annual estimated mortality of 100
penguins (Tamini et al. 2002), and in the mid-water
coastal fishery targeting silversides Odontesthes spp.
along the northern coast of Argentina (3 captures
across 28 d; Seco Pon et al. 2013).
Small-scale fisheries (gillnets). Very few data are
available from small-scale fisheries, including coastal
gillnets, from areas that overlap with Magellanic
penguins at sea. This is an important knowledge gap
given the high risk to penguins from gillnets. Bycatch
has been recorded in the south of Argentina: from
1995−1999, around 1000 penguins were estimated to
have been killed annually (Schiavini et al. 2005, P.
Gandini & E. Frere unpubl. data).
Falkland Islands (Islas Malvinas)
Trawl fisheries. The Doryteuthis gahi squid fishery
in the Falkland Islands comprises 16 bottom trawlers
operating to the east and southeast of the islands,
376
1A dispute exists between the Governments of Argentina and
the UK concerning sovereignty over the Falkland Islands (Is-
las Malvinas), South Georgia and the South Sandwich Is-
lands (Islas Georgias del Sur y Islas Sandwich del Sur) and
the surrounding maritime areas.

Crawford et al.: Penguin bycatch in global fisheries 377
Species
Argentina
Falklands/Islas Malvinas
Uruguay
Brazil
Chile
Peru
Ecuador
South Africa
Namibia
New Zealand
Australia
Antarctica & Southern
Ocean Islands
(a) Gillnet bycatch
Magellanic
Galápagos
Humboldt
African
Little
Southern
rockhopper
Northern
rockhopper
Macaroni
Royal
Snares
Fiordland
Erect-crested
Yellow-eyed
Chinstrap
Gentoo
King
Emperor
Adelie
Table 1. Relative susceptibility of penguin species to bycatch by gear type: (a) gillnet, (b) trawl, (c) longline, (d) purse seine. Key to relative impact ranks: red = high (mor-
tality recorded in several studies), yellow = medium (mortality recorded at a high level in 1 study, or a low level in more than 1 study), green = low (no mortality recorded,
or isolated cases), grey = not applicable (species not present in this area), black = unknown (potential for impact, but limited studies for this gear type/species/region)
Species
Argentina
Falklands/Islas Malvinas
Uruguay
Brazil
Chile
Peru
Ecuador
South Africa
Namibia
New Zealand
Australia
Antarctica & Southern
Ocean Islands
(b) Trawl bycatch
Magellanic
Galápagos
Humboldt
African
Little
Southern
rockhopper
Northern
rockhopper
Macaroni
Royal
Snares
Fiordland
Erect-crested
Yellow-eyed
Chinstrap
Gentoo
King
Emperor
Adelie

Endang Species Res 34: 373– 396, 2017
378
Species
Argentina
Falklands/Islas Malvinas
Uruguay
Brazil
Chile
Peru
Ecuador
South Africa
Namibia
New Zealand
Australia
Antarctica & Southern
Ocean Islands
(c) Longline bycatch
Magellanic
Galápagos
Humboldt
African
Little
Southern
rockhopper
Northern
rockho
pp
er
Macaroni
Royal
Snares
Fiordland
Erect-crested
Yellow-eyed
Chinstrap
Gentoo
King
Emperor
Adelie
Table 1 (continued)
Species
Argentina
Falklands/Islas Malvinas
Uruguay
Brazil
Chile
Peru
Ecuador
South Africa
Namibia
New Zealand
Australia
Antarctica & Southern
Ocean Islands
(d) Purse seine bycatch
Magellanic
Galápagos
Humboldt
African
Little
Southern
rockhopper
Northern
rockho
pp
er
Macaroni
Royal
Snares
Fiordland
Erect-crested
Yellow-eyed
Chinstrap
Gentoo
King
Emperor
Adelie

Crawford et al.: Penguin bycatch in global fisheries
largely within 65 km of the coast. Between 1995 and
2001, low levels of net-related bycatch were recorded
for several penguin species, including 1 king pen-
guin, 6 gentoo penguins and 7 Magellanic penguins,
most of which occurred off Volunteer Point (51° S,
57° W) in an area now closed to fishing (Falkland
Islands Fisheries Department unpubl. data).
The mixed finfish and skate fleet mainly fishes in
deeper water (>140 m), so while this fishery does
cause some seabird mortalities (primarily black-
browed albatrosses Thalassarche melanophris), pen-
guin fatalities are believed to be very rare: a single
king penguin mortality was recorded in 2009 (Falk-
land Islands Fisheries Department unpubl. data).
In October 2012, a research trip on a commercial
bottom trawler caught 23 gentoo penguins over 3 d to
the north-east of the islands: 6 of these birds drowned,
and 17 survived and were released (Roux et al. 2012).
Dissection revealed that several penguins had been
feeding on fishery discards. Since 1988, observers
have been placed regularly on comparable vessels
fishing in the same area, and penguin captures have
not been reported (J. Pompert pers. obs.).
Jigging fisheries. Squid jigging (targeting Illex
argentinus) accounts for the largest component of
catches in the Falkland Islands, with vessels operat-
ing between the Falkland Islands and Argentina.
Barton (2002) reported that low numbers of pen-
guins are occasionally hooked by the small barbless
hooks on the jigging lures and brought on board.
After a period on the vessel they are released ap -
parently un harmed. Observers present on the ship
inspect animals for cuts, but have generally reported
injury to be of a very minor nature (J. Pompert pers.
obs.).
Longline fisheries. The single Patagonian toothfish
Dissostichus eleginoides demersal longline vessel
fishing in Falkland Islands waters is Marine Steward-
ship Council certified and has not recorded any pen-
guin bycatch (with high levels of observer coverage:
40−80% of effort was observed between 2002 and
2012; Andrews et al. 2014). In September 2006, an
ex perimental longline fishery for kingclip Genypterus
blacodes hooked and drowned 5 gentoo penguins
(Falkland Islands Fisheries Department unpubl. data).
Despite 2 significant bycatch events, overall pen-
guin bycatch is believed to be rare in Falkland
Islands fisheries. Gentoo penguins appear to be the
most susceptible species, although Magellanic and
southern rockhopper penguins that breed in the
Falklands may be vulnerable on their wintering
grounds in Argentinean, Uruguayan and Brazilian
waters.
Uruguay
Minimal penguin bycatch data are available for
Uruguay, though there is some information in the
National Plan of Action–Seabirds (Domingo et al.
2015).
Trawl fisheries. The demersal trawl fishery for
Argentine hake has recorded Magellanic penguins
entangled in nets, but there is no quantitative esti-
mate of the numbers of penguins involved. The
coastal trawl fishery targeting whitemouth croaker
(locally known as corvina) Micropogonias furnieri
and weakfish Cynoscion guatucupa uses 2 vessels
trawling a single net, and incidental captures of
Magellanic penguins have been reported (Karumbé
unpubl. data).
Gillnet fisheries. Both artisanal and industrial gill-
net fisheries operate in Uruguay, using bottom and
surface-set nets, though primarily the former. The
main target species are whitemouth croaker and
weakfish, and seabird bycatch has been detected,
but has not been recorded to species level. In line
with reports from Brazil, it seems likely that it is Mag-
ellanic penguins that are captured in these fisheries.
Purse seine fisheries. National observers have re -
corded Magellanic penguin bycatch in the Uruguayan
purse seine fishery targeting anchoita Engraulis an -
choita, but no quantitative data are available.
Brazil
Gillnet fisheries. Gillnets are the only fishing gear
in which substantial penguin bycatch has been re -
corded in Brazil, and those operating in the south,
overlapping with Magellanic penguins, can be
broadly placed in 3 categories: the surface or driftnet
fishery targeting bluefish Pomatomus saltatrix; bot-
tom-set gillnets targeting weakfish and Argentine
croaker Umbrina canosai; and bottom-set gillnets tar-
geting whitemouth croaker. Penguin bycatch has
been recorded in all of these fisheries, but vessels tar-
geting bluefish are of greatest concern (L. Bugoni
unpubl. data).
Cardoso et al. (2011) undertook observations on a
gillnet vessel typical of the Passo de Torres fleet in
southern Brazil, which deployed both driftnets for
bluefish and bottom-set gillnets for weakfish and
Brazilian codling Urophicys brasiliensis. From 17 ob -
served sets over 8 d, 68 Magellanic penguins (pre-
dominantly adults) were killed: 56 in driftnets (mean
± SD: 2.65 ± 1.47 birds km−1 of net d−1), and 12 in bot-
tom-set gillnets (0.22 ± 0.16 birds km−1 of net d−1).
379

Endang Species Res 34: 373– 396, 2017
Based on interviews with fishermen, Vasconcellos
et al. (2014) reported that incidental capture of pen-
guins is more common in both drift and bottom-set
gillnets near the Uruguayan border. This is consistent
with the distribution of larger schools of anchoita, the
main Magellanic penguin prey north of 47° S (Frere
et al. 1996). In 2008, 280 gillnet vessels were fishing
with driftnets and/or bottom-set gillnets from ports in
southern Brazil (IBAMA/CEPERG 2009, UNIVALI
2009). A recent estimate suggested that over 350 ves-
sels operated in Santa Catarina and Rio Grande do
Sul states (Vasconcellos et al. 2014), so the scale of
bycatch over this entire fleet could be substantial.
Chile
Gillnet fisheries. Based on observation of fishing
activities, beached bird surveys and interviews with
fishermen, 605 Humboldt and 58 Magellanic pen-
guins were reported drowned as bycatch between
1991 and 1996 in the corvina Cilus gilberti gillnet
fishery operating in central Chile (~33° S; Simeone et
al. 1999).
Mass penguin mortality events of several hundred
birds in south-central Chile (39° S) have also been
linked to bycatch. Schlatter et al. (2009) reported
1380 stranded Magellanic penguins (adults and juve-
niles) at Queule. Based on injuries consistent with
entanglement and on interviews with local fisher-
men, these authors attributed this mortality event to
drowning in artisanal gillnets targeting southern rays
bream Brama australis and corvina. Skewgar et al.
(2009), Pütz et al. (2011) and Suazo et al. (2013) also
reported a low number of records of entangled Mag-
ellanic penguins in local gillnet fisheries close to
Chiloé Island (42° S, 74° W) and Chonos Archipelago
(45° S, 73° W) in southern Chile.
Purse seine fisheries. Recent observations in both
artisanal and industrial purse seine vessels targeting
sardine Strangomera bentincki show that penguins
can be captured in this gear: a total of 4 Magellanic
penguins were recorded in 16 monitored trips in
2014 (Albatross Task Force-Chile unpubl. data),
which merits further investigation.
Longline fisheries. There are scattered records of
penguin bycatch from the sub-Antarctic fjords in
southern Chile. Among these are 3 southern rock-
hopper penguins from 17 monitored trips (J. Ojeda
pers. comm.), which were hooked in small-scale ver-
tical drifting longlines set for southern hake Merluc-
cius australis (see Moreno et al. 2006, Suazo et al.
2013).
Overall, it is thought that small-scale fisheries, par-
ticularly gillnets and purse seines, are the main fish-
eries threatening penguins in Chile (Suazo et al.
2014).
Peru
Evidence of incidental catch and mortality of pen-
guins in Peruvian gillnet fisheries has been noted
since the early 1980s (Duffy et al. 1984, Hays 1984,
Araya 1988).
Gillnet fisheries. Incidental capture in gillnets has
previously been reported as the main cause of mor-
tality for the Punta San Juan (12° S, 77° W) and
nearby colonies, with about 400 Humboldt penguins
reported dead in 1992 (Zavalaga & Paredes 1997).
Observers stationed at San Juan port recorded land-
ings of 922 dead Humboldt penguins between 1991
and 1998 (Majluf et al. 2002). The majority of these
were captured in the surface driftnet fleet targeting
cojinova Seriolella violacea, with lower bycatch rates
in fixed demersal gillnets (Majluf et al. 2002). This
study also highlighted that driftnets were usually set
through the night, and penguins were thought to
have been captured when staying at sea overnight,
resting on the surface (Majluf et al. 2002). It is sus-
pected that mortality could be exacerbated by reten-
tion of incidentally captured penguins for human
consumption, as active targeting of penguins with
gillnets has been recorded (P. McGill pers. comm.).
Humboldt penguin bycatch has also been recorded
in the demersal set net fishery for guitarfish Rhino-
batos planiceps in Constante (5° S, 99° W) and in the
driftnet fishery for sharks and rays in Salaverry (8° S,
79° W; Alfaro-Shigueto et al. 2010). Between 2000
and 2015, this observer programme recorded bycatch
of 22 penguins from 6 fishing ports, the majority of
which were captured in surface driftnets set at night
(Pro Delphinus unpubl. data).
Artisanal vessels, many of which use gillnets, are
the main threat to penguins. Considering that there
are more than 10 000 such vessels in the country
(Goya et al. 2011), the impact of this large fleet may
be significant for Humboldt penguins.
Ecuador
The Galápagos penguin S. mendiculus is an En -
dangered species endemic to the Galápagos Islands,
with an estimated population of 1200 individuals
(BirdLife International 2017a), and is the only pen-
380

Crawford et al.: Penguin bycatch in global fisheries
guin species breeding in Ecuador (Harris 1973). With
the creation of the Galápagos Marine Reserve (GMR),
industrial fishing was banned and fishing rights were
granted exclusively to the local small-scale, or arti-
sanal, fishing sector (MAE-DPNG 2006, Asamblea
Nacional 2015).
Despite being illegal in the GMR, gillnets are reg-
ularly used in shallow areas to target schooling fish.
Penguin bycatch has been recorded in these nets in
the waters off Villamil, Isabela Island, but quantita-
tive figures are not available (A. Steinfurth pers. obs.,
Galápagos National Park Directorate unpubl. data).
A small number of birds have also been found entan-
gled in discarded ‘ghost’ fishing gear in the Bolivar
Channel (potentially originating outside the GMR;
Galápagos National Park Directorate pers. obs.).
With few records of fisheries-related penguin mor-
tality in Ecuador, and no legal gillnet fishery, penguin
bycatch does not appear to be a major threat. How-
ever, it is suspected that bycatch is under-reported,
particularly because it occurs in illegal fisheries.
There is a clear need for contemporary data to ascer-
tain the threat illegal fishing poses to the endangered
Galápagos penguin.
Southern Africa
The African penguin S. demersus is endemic to
southern Africa and is the only breeding penguin on
the African continent (Crawford et al. 2011). Although
vagrant birds have been found as far north as Gabon
on the African west coast and Mozambique on the
east coast, this species breeds only from central
Namibia to Bird Island, Algoa Bay, in South Africa’s
Eastern Cape (Crawford et al. 2013). Some birds, par-
ticularly non-breeding birds (Shelton et al. 1984) and
juveniles (Sherley et al. 2017) do venture north into
Angolan waters.
The species is currently listed as Endangered
(BirdLife International 2017a) due to a large popula-
tion decrease over the preceding 30 yr (Crawford et
al. 2011). The main cause of the decrease is thought
to be related to prey (sardine Sardinops sagax and
anchovy Engraulis encrasicolus) availability (Craw-
ford et al. 2011). Mortality of African penguins in fish-
ing gear is unquantified in the region (Figs. 1 & 2).
South Africa
Purse seine fisheries. South Africa’s purse seine
fishery targets mainly sardine, anchovy and red eye
round herring Etrumeus whiteheadi and is the coun-
try’s largest commercial fishery by landed mass
(Prochazka 2014). It is thought that African penguins
avoid fishing vessels, although there are conflicting
accounts (H. Terblanche and S. Hampton pers.
comm.). There have been no reports of African pen-
guin captures in these nets (L. Pichegru pers. comm.),
but there are no independent observers in this fish-
ery to verify this.
Gillnet fisheries. Beach seine and gillnet fisheries
are active throughout South Africa (Prochazka 2014),
with the largest targeting harder (mullet) Liza richard-
sonii. These fisheries are effort managed, but there
are substantial numbers of illegal fishers (Prochazka
2014). Much of the effort is focussed around river
mouths, estuaries and on the west coast, where the
fisheries overlap with the distribution of penguin
breeding colonies.
Several hundred African penguins were reportedly
caught in gillnets around Dassen and Robben Islands
until gillnet exclusion zones were put in place in
2001. Any bycatch that now occurs in these areas
would be from illegal or ghost gillnets. St. Joseph
shark Callorhinchus capensis gillnets, and those tar-
geting smooth-hound sharks Mustelus mustelus and
various species of ‘linefish’ (those generally caught
on lines from shore or small vessels), are likely to be
responsible for penguin bycatch, if any occurs.
Bycatch from illegal fishing is exceedingly difficult to
quantify (all information from S. Lamberth pers.
comm.).
Trawl fisheries. Three main trawl fisheries operate
in South Africa: the demersal trawl fishery targeting
shallow-water cape hake Merluccius capensis and
deep-water cape hake M. paradoxus, the inshore
trawl fishery targeting a mix of species including
Agulhas sole Austroglossus pectoralis and the cape
hakes, and the midwater trawl fishery targeting cape
horse mackerel Trachurus capensis. The demersal
trawl fishery is unlikely to be a source of bycatch
mortality as it has minimal overlap with foraging
African penguins, and the species is not known to
take discards. Fisheries observers on trawl vessels
(approximately 20% observer coverage) have never
recorded penguin mortality in this fishery (B. Maree
pers. comm.).
No instances of penguin mortality have been
recorded in the inshore trawl fishery. African pen-
guins do not forage on the species targeted by the
fishing vessels and are therefore unlikely to be in the
same area at the time of fishing.
Longline fisheries. Limited longline fishing for tuna
and billfish occurs in South African and Namibian
381

Endang Species Res 34: 373– 396, 2017
waters. Lines are generally set offshore (B. Lebepe
pers. comm.), beyond the average foraging range of
the African penguin, which is around 20 km from the
coast (Wilson et al. 1988, Ludynia et al. 2012).
Namibia
Namibia has purse seine and longline fisheries
similar to South Africa, and the issues around pen-
guin bycatch (or lack thereof) are currently believed
to be similar and are not discussed further. However,
while no penguin mortalities have been recorded, it
should be emphasised that there is limited observer
coverage.
Gillnet fisheries. The use of gillnets is prohibited in
Namibia, but the small-scale artisanal/recreational
use of set nets is tolerated by fisheries inspectors in a
bay close to the African penguin colony on Halifax
Island. Small pieces of broken gillnet are regularly
found washed up on beaches in the area (J. Kemper
pers. obs.). Although this is a very localized and small-
scale fishery (perhaps fewer than 8 rowing boats), the
close proximity of gillnet fishing activities (and the
projected path of ‘ghost’ net fragments drifting with
the prevailing current) to Halifax Island (26° S, 15°E)
and the overlap with penguin foraging paths consti-
tute a potential threat. Gillnet entanglement has been
directly observed for crowned cormorants Microcarbo
coronatus, and fragments of gillnets have been re -
covered from crowned and cape cormorant Phalacro-
corax capensis nests (J. Kemper pers. obs.).
Trawl fisheries. Horse mackerel catches in the mid-
water trawl industry have dominated Namibian land-
ings for the last 30 yr (Roux et al. 2013). While no
African penguin bycatch has been reported (J. Kem-
per pers. obs.), this is an important knowledge gap to
fill due to the scale of the fishery.
Australasia
Nine penguin species are found in Australia, New
Zealand and their sub-Antarctic territories. This sec-
tion excludes the islands Macquarie, Heard, McDonald
(to Australia) and Antipodes, Auckland and Campbell
(to New Zealand), which are considered below in the
‘Southern Ocean Islands’ section.
The focus for this region is on 3 of the 4 New
Zealand endemics: the Endangered yellow-eyed
Mega dyptes antipodes, the Vulnerable Snares Eu -
dyptes robustus and Fiordland E. pachyrhynchus
penguins, and the little penguin Eudyptula minor
(BirdLife International 2017a). Little penguins are
the only mainland species in Australia; they are also
found on the New Zealand mainland. Recent research
suggests that Australian and New Zealand little
penguins are separate species (Grosser et al. 2015).
This may have implications for conservation status
and actions, but since the little penguin is formally
recognised as a single species at the time of writing
(BirdLife International 2017b), we consider it here
as a single species.
Yellow-eyed penguins also have populations on
the sub-Antarctic Auckland and Campbell Islands
(Seddon et al. 2013), but these birds are genetically
isolated from the mainland population (Boessenkool
et al. 2009) and should be considered separately for
management purposes (Ellenberg & Mattern 2012).
Both the Australian and New Zealand commercial
fishing fleets include larger industrial vessels operat-
ing offshore, primarily utilising trawls and longlines,
in addition to smaller inshore operations that use a
variety of gear types. Of these, gillnets are believed
to be of greatest concern for penguins, although in -
dependent observer data are sparse (Table 1).
New Zealand
Gillnet fisheries. Gillnets, also called set nets in
New Zealand, are the primary gear type of concern
with regard to penguin bycatch. The recreational
sector focusses effort inshore, whereas the commer-
cial sector fishes more extensively on the continental
shelf around the New Zealand mainland (Ministry of
Primary Industries 2015). The number of gillnet ves-
sels has decreased in recent years; at present, around
330 commercial vessels use this gear in New Zealand
waters (Ministry of Primary Industries 2012a), with
operations primarily targeting demersal fish species
(such as tarakihi Nemadactylus macropterus, jock
stewart Helicolenus percoides, rig Mustelus lenticu-
latus and school shark Galeorhinus galeus). These
fisheries overlap spatially with the benthic foraging
yellow-eyed penguin.
In 2008, an inshore gillnet ban of between 2 and 4
nautical miles offshore was introduced around the
South Island to reduce Hector’s dolphin Ce pha lo -
rhynchus hectori bycatch. This ban may have con-
tributed to a reduction in the impact of recreational
gillnets on penguins, though is unlikely to have sub-
stantially reduced the risk from commercial gillnets,
as key yellow-eyed penguin foraging grounds have
been identified 15−20 km from the shore (Mattern et
al. 2007, Ellenberg & Mattern 2012).
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Crawford et al.: Penguin bycatch in global fisheries
Darby & Dawson (2000) reported a total of 72 en -
tanglements of yellow-eyed penguins (62% of which
were adults), corresponding to ~12 mortalities yr−1,
primarily around the Otago Peninsula and north
Otago (45° S, 171°E) between 1979 and 1997. Bycatch
in commercial gillnets occurred predominantly in
bottom set nets at depths >70 m, more than 15 km
offshore.
Hocken (2005) analysed the cause of death in 114
yellow-eyed penguins collected opportunistically from
1996−2003 and found that at least 6 had been trapped
in nets, diagnosed by characteristic bruising about the
neck and shoulders. Anecdotal evidence suggests
that bycatch continues to occur, and that previously
reported numbers may represent an underestimate
(Ellenberg & Mattern 2012).
More recent observer coverage of commercial
gillnetters reported 9 captures of yellow-eyed pen-
guins over a 5 yr period (2005−2010; Rowe 2008,
2009, 2010, Ramm 2010, 2012a,b), much lower than
Darby & Dawson (2000), which itself was considered
to be an underestimate of actual bycatch. The low
number of yellow-eyed penguin deaths reported
through the observer programme in recent years
may be related to overall low levels of observer cov-
erage (1.5% of effort), and a lack of monitoring on
gillnetters operating in important yellow-eyed pen-
guin foraging areas (Richard & Abraham 2013). Based
on the limited ob server dataset, Richard & Abraham
(2015) estimated fatalities of mainland population
yellow-eyed penguins in gillnets to be on average
35 birds yr−1 (range: 16−60 birds), with the largest
proportion (ca. 70%) taken in demersal nets set for
sharks. Population viability analysis shows that this
level of mortality could have population-level effects
(McKinlay 1997). In 2016, another 4 yellow-eyed
penguins were recorded as bycatch in commercial
gillnetters targeting school shark (ob server coverage
2.8% effort; Ministry of Primary Industries 2017).
Two of these birds were caught off the Waitaki River
Mouth, 1 off Kaka Point and another off Stewart
Island. Given the species’ precarious situation, with
low numbers (216 breeding pairs on the New Zealand
South Island in 2015−16) and an overall decreasing
population at previous strongholds (76% decline
be tween 1996 and 2016; Ellenberg & Mattern 2012,
Mattern et al. 2017), the effects of gillnet bycatch on
the species may be significant and merits closer
examination.
Little information is available to assess the extent to
which other penguin species might be affected by
gillnetting around the New Zealand mainland. Darby
& Dawson (2000) noted that the Otago Museum holds
9 records of little penguins taken in gillnets. Despite
considerable overlap of inshore fisheries and impor-
tant penguin foraging grounds, the observer pro-
gramme had not reported any incidents of little pen-
guin bycatch prior to 2016 (Richard & Abraham 2013),
probably due to extremely low observer coverage on
gillnetters operating in penguin foraging areas. Efforts
to increase independent observer coverage to 2.8%
in 2016 saw 8 little penguins bycaught in a single
observed net set for butterfish Odax pullus across a
channel in the Stewart−Snares area (Ministry of Pri-
mary Industries 2017).
The inshore gillnet ban in southern New Zealand
may benefit little penguins in those areas, although
exemptions have been granted for net fishing adja-
cent to key penguin breeding sites (e.g. Motuara
Island; Numata et al. 2000, Mattern 2001, Ministry of
Primary Industries 2012b, 2015). Most of the rest of
New Zealand, where little penguins are widely dis-
tributed, has few limits on setting nets.
The rugged and exposed coastlines along most of
the Fiordland penguin breeding range largely pre-
vent gillnetting operations (Mattern 2013a). How-
ever, many Fiordland penguins forage sympatrically
with yellow-eyed penguins in areas where commer-
cial gillnetters operate, especially in the western
Foveaux Strait. While only 3 bycatch incidents have
officially been reported for this species (Ramm 2010,
Ministry of Primary Industries 2017), limited ob server
coverage makes it difficult to undertake a meaning-
ful assessment of the level of threat.
Snares penguins only breed on the Snares Islands,
200 km south of the mainland, and are therefore less
exposed to gillnetting. However, birds forage to the
south of Stewart Island during chick-rearing (Mat-
tern et al. 2009, Mattern 2013b), an area fished by
gillnets. The risk of gillnet bycatch for Snares pen-
guins is likely to be lower than for other mainland
species, but not negligible.
Australia
Gillnet fisheries. The majority of reported penguin
bycatch incidents in mainland Australia have oc -
curred in gillnets, but data are sparse, with few ob -
server data, making assessment of the bycatch risk to
little penguins in Australia difficult. Nevertheless,
there are published records of little penguins being
entangled and drowned in active or discarded gill-
nets in Australia, including Victoria (Norman 2000),
South Australia (Copley 1996, Knight & Vainickis 2011)
and Tasmania (Stevenson & Woehler 2007). More sub-
383

Endang Species Res 34: 373– 396, 2017
stantial bycatch events have been recorded, includ-
ing some 50 dead, entangled little penguins washed
ashore on Chalky Island (Bass Strait, Tasmania) in
2015 (Tasmanian Department of Primary Industries,
Parks, Water and Environment unpubl. data). A num-
ber of other bycatch events involving little penguins
have been reported from southeast Tasmania (E. J.
Woehler unpubl. data).
Most data on bycatch of little penguins comes from
around Phillip Island, southeast Australia (38° S,
145° E), one of the largest colonies of this species
(Nisbet & Dann 2009, Dann & Chambers 2013). The
majority of little penguins breeding at Phillip Island
feed within 15 km of the coast in northern Bass Strait,
where there is considerable overlap with gillnets, or
in Port Phillip Bay (Collins et al. 1999, Chiaradia et al.
2007, McCutcheon et al. 2011, Pelletier et al. 2014).
Most commercial nets are set at between 40 and
120 m depth (Kailola et al. 1993, Bulman et al. 2006),
overlapping with the foraging depths of little pen-
guins that can dive to a max dive depth of 72 m but
normally forage at 20−40 m (Ropert-Coudert et al.
2006, 2009). Recreational gillnetters, where permitted,
fish closer inshore than commercial vessels, increas-
ing the risk of interaction, particularly around colonies.
In 2007, 23 penguins were caught in a shark net
15 km from Queenscliff, Victoria (Phillip Island Nat -
ure Parks, unpubl. data), and 25 dead birds were
found on Altona Beach in Port Phillip Bay, Victoria, in
2014. Necropsies of the latter concluded that the
birds had died after becoming entangled in a gillnet
(O’Doherty 2014).
There are 42 records of dead flipper-banded little
penguins from Phillip Island reported as ‘bird tan-
gled in fishing gear’ to the Australian Bird and Bat
Banding Schemes (ABBBS) or to the Phillip Island
Nature Parks. The relative proportions killed in gill-
nets or through entanglement in fishing line/snared
by hooks are unknown, but both types of mortality
have been reported (ABBBS and Research Depart-
ment, Phillip Island Nature Parks unpubl. data).
In Tasmania, where approximately 10 000 gillnets
are registered (plus an unknown number of ‘many
more’ unregistered nets), a study undertaking research
on gillnets resulted in the bycatch of 5 little penguins,
despite the study being designed carefully to avoid
seabird interactions. During a recent survey of re -
creational gillnetters, over a quarter of respondents
acknowledged they experienced interaction with sea -
birds during gillnetting; of these, 39% reported en -
tanglements of penguins (Lyle et al. 2014). This indi-
cates that bycatch levels may be higher than those
reported to date.
While spatial closures have been implemented
around some penguin colonies in Tasmania (Lyle et
al. 2014), there remain many colonies without such
protection. Improving observation of gillnet fisheries
to identify bycatch hotspots and inform protected
area identification for little penguins, particularly
around Tasmania, are thus priority actions.
Antarctica and the Southern Ocean Islands
including the CCAMLR area
The Antarctic and the Southern Ocean islands
host, by far, the largest number of penguins com-
pared with the other regions in this review (García-
Borboroglu & Boersma 2013). The region also has the
lowest human population and an absence of recre-
ational fishing beyond occasional research efforts at
national research facilities. The majority of the fishing
effort is larger-scale, particularly trawling, long lining
and squid jigging, primarily under the management
of the CCAMLR southward of the approximate loca-
tion of the Antarctic Polar Front. However, this is not
the case for all parts of this region, and illegal, unre-
ported and unregulated (IUU) fishing remains a con-
cern for some CCAMLR and non-CCAMLR areas of
the Southern Ocean.
Here, we consider penguin bycatch within the
CCAMLR area as a single ‘unit’ (given the broadly
uniform fisheries management requirements), with
separate assessments for penguin populations out-
side the Convention area: the Tristan da Cunha group,
Amsterdam Island, St. Paul Island, the New Zealand
sub-Antarctic islands (Antipodes, Auckland and
Campbell) and Macquarie Island.
Tristan da Cunha and Gough Island
The Tristan da Cunha group of islands in the South
Atlantic, a UK Overseas Territory, hosts the vast
majority of the world’s breeding northern rockhop-
per penguins Eudyptes moseleyi (~85% of the global
population estimate of 240 300 breeding pairs; Cuth-
bert et al. 2009, Robson et al. 2011, Tristan da Cunha
Government and Royal Society for the Protection of
Birds unpubl. data).
Longline fisheries. During 2015 and 2016, a single
vessel fished for tuna (Thunnus spp.) around Tristan
using pelagic longlines. With such a low level of effort,
it is believed to have no impact on the northern rock-
hopper population, although there are no data avail-
able on bycatch from this vessel (J. Glass pers. obs.).
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Crawford et al.: Penguin bycatch in global fisheries
Gillnet fisheries. Bycatch of northern rockhopper
penguins in driftnets was recorded from the Tristan
exclusive economic zone (EEZ) in the late 1980s/
early 1990s (Ryan & Cooper 1991). Reports from re -
search vessels, crew members and evidence from a
vessel that ran aground in South Africa in 1990 sug-
gest that mortalities from the estimated 160 vessels
that operated in the area could have been substantial
(Ryan & Cooper 1991). The 1992 UN Resolution ban-
ning high seas driftnets means that legal gillnet fish-
eries are no longer of concern, though IUU fishing is
suspected.
Trawl fisheries and IUU activity. After an exploratory
seamount trawl fishery in Tristan da Cunha waters,
which operated for 14 d in 2015, a single commercial
trawler targeted splendid alfosino Beryx splendens,
but mainly caught rosefish Helicolenus mouchezi
and cape redfish Sebastes capensis (J. Glass pers.
obs.).
No penguins were reported caught, but 6 different
types of fishing gear were hauled aboard by the
trawler, including jigging gear, gillnets, trawl nets
and longlines. Although there are no guarantees as
to the provenance of these gears, they were thought
to have been used recently based on their condition,
and were suspected to have been previously de -
ployed illegally within the Tristan EEZ (J. Glass pers.
obs.).
The illegal use of gillnets and trawls is therefore
the primary bycatch-related concern for northern
rockhopper penguins on Tristan da Cunha, but there
is little capacity to assess IUU activity and enforce
fisheries regulations for this small group of islands
with a large EEZ. In addition, the wintering move-
ments of northern rockhopper penguins (and their
overlap with fisheries) are not fully understood.
Amsterdam Island and St. Paul Island
These 2 islands are French Southern Territories in
the southern Indian Ocean and host the remainder of
the northern rockhopper penguin population (~15 %;
Guinard et al. 1998, Cuthbert 2013, BirdLife Interna-
tional 2017). A St. Paul rock lobster Jasus paulensis
fishery is active around the islands, but this is not
presently considered to be a threat to penguins (how-
ever, historically penguins were used as bait in lob-
ster pots, with suspected population-level effects;
Guinard et al. 1998). There is also overlap with long-
line vessels fishing under Indian Ocean Tuna Com-
mission jurisdiction, but little information is available
on bycatch from these vessels (TAAF 2011).
Antipodes, Auckland and Campbell Islands
New Zealand’s sub-Antarctic islands host 3 breeding
penguin species: yellow-eyed, southern rockhopper
E. chrysocome and erect-crested E. sclateri penguins.
Longline and trawl fisheries. Fishing practices in
the sub-Antarctic region are principally limited to
longlining or trawling. While observer programmes
have recorded significant mortality of flying seabirds
in New Zealand trawl and longline fisheries in recent
years (Ramm 2010, 2012a,b, Dragonfly Data Science
2016), cases of penguin bycatch are practically
unknown. One erect-crested penguin was foul hooked
in the flipper by a ling (Molva spp.) longliner during
hauling, and was subsequently released alive (Ramm
2010).
Without any indication of interactions, the number
of estimated fisheries-related fatalities for all sub-
Antarctic penguin species currently stands at 0 for
these New Zealand islands (Richard & Abraham
2013), though ob server coverage is variable (broadly
lower in inshore trawl compared to offshore fisheries;
Dragonfly Data Science 2016).
Macquarie Island
Macquarie Island holds breeding populations of
southern rockhopper, gentoo, king and royal pen-
guins. No recreational gillnetting or trawling is per-
mitted (AFMA 2016a,b).
Longline fisheries. A Patagonian toothfish fishery
uses longlines around Macquarie Island (AFMA
2016a). As of the end of 2012, no penguins have been
reported as bycatch.
CCAMLR Convention Area
The CCAMLR Convention Area supports an over-
whelming majority of the world’s penguins (García-
Borboroglu & Boersma 2013). The area is managed
under a regulatory framework that requires fisheries
not to expand beyond the limits of the available infor-
mation on their impacts (CCAMLR 2015a). A number
of conservation measures are in place, including spe-
cific measures to mitigate against seabird mortality
(including seasonal closures and gear restrictions),
several marine protected areas (MPAs) and rigorous
bycatch reporting procedures (CCAMLR 2015a).
Four main species are targeted across the entire
convention area: Patagonian toothfish and Antarctic
toothfish Dissostichus mawsoni (13 licensed fisheries,
385

Endang Species Res 34: 373– 396, 2017
largely demersal longliners), mackerel icefish Cham -
psocephalus gunnari (caught using midwater and
bottom trawls) and Antarctic krill Euphausia superba
(caught using midwater trawls and beam trawls)
(CCAMLR 2015a).
There are high levels of observer coverage across
the area; icefish and toothfish fisheries have a re -
quirement for 100% coverage, and target coverage
of 50% in the krill fishery. This makes the Convention
Area data-rich compared to other regions re viewed
here, and gives high confidence that the bycatch
events highlighted below are representative of fleet-
level impacts. Broadly, penguin bycatch is a rare
occurrence in the legal CCAMLR fisheries. However,
IUU fishing remains a concern in some regions, and
is therefore considered separately below. Data for
this section were mostly retrieved from CCAMLR
Fisheries Reports (available on the CCAMLR web-
site), though some additional, detailed information is
included for South Georgia and the South Sandwich
Islands.
Longline fisheries. Isolated incidents of penguin
bycatch in toothfish fisheries have been recorded.
Chilean industrial longliners for Patagonian toothfish
recorded penguin bycatch events in CCAMLR Sub-
area 48.3 (South Georgia) during 1995 (Moreno et
al. 1996), with both gentoo and macaroni penguins
hooked and released alive. Similarly, a gentoo pen-
guin was captured and released in the longline fish-
eries around the Kerguelen Islands between 1994
and 1997 (Weimerskirch et al. 2000). In the 1997/98
fishing season, 5 macaroni penguins were foul hooked
in the body around the Prince Edward Islands (a
South African territory) (Nel et al. 2002). From 2001
to 2003, 1 king, 1 gentoo, 2 macaroni and 1 unidenti-
fied penguin were among the anecdotal catch from
around Crozet and Kerguelen Islands (French South-
ern Territories) (CCAMLR 2004, Delord et al. 2005).
A single southern rockhopper mortality was recorded
in the same area in both the 2005/06 (CCAMLR 2006)
and the 2010/11 seasons (CCAMLR 2011).
In 2005, a gentoo penguin was caught in a demer-
sal longline in the South Georgia and South Sand-
wich Islands Maritime Zone, although on this occa-
sion it was entangled in the line. In 2009, a chinstrap
penguin Pygoscelis antarctica was injured in the
exploratory toothfish fishery around the South Sand-
wich Islands (CCAMLR 2013). A single penguin (spe-
cies not specified) was hooked in the foot in 2014, but
this bird was released alive and in good condition (K.
Ross pers comm.). No other penguin bycatch inci-
dents were observed in this area between 2001 and
2014.
The most recent recorded mortality comes from the
toothfish fishery around Heard Island (Australian EEZ)
in 2014, where 1 southern rockhopper was killed
(CCAMLR 2014).
Trawl fisheries. The only penguin bycatch recorded
in CCAMLR mackerel icefish trawl fisheries was in
2008, when 2 king penguins were caught around
South Georgia. There are only 2 active mackerel ice-
fish fisheries in the region: the South Georgia fishery
(total allowable catch of 2074 t in 2016/17), and a
smaller fishery around Heard Island (total allowable
catch of 482 t in 2015/16 season; CCAMLR 2016a).
The last record of penguin (species unknown) by -
catch in krill trawlers was in 2007, but it is not known
whether this interaction occurred during active fish-
ing or on retrieval of the gear. Although observer cov-
erage in this fishery is lower than in other CCAMLR
fisheries, it is not suspected that penguin bycatch is
an issue. Postulated reasons for the low penguin by -
catch in this fishery have centred on the slow trawl-
ing speeds (2−3 knots) and the potential for some seal
bycatch mitigation devices to allow penguins to
escape (K. Reid pers. comm.).
IUU fisheries. Although IUU fishing is believed to
have decreased markedly since the 1990s, when ille-
gal catch was estimated to be 6 times that of legal
catch in the Convention Area (CCAMLR 2016b), it
continues to occur in some CCAMLR regions. It can
be safely assumed that these vessels will not report
by catch and regularly use gear types that are not
permitted in the CCAMLR area, particularly gillnets,
which pose a significant threat to penguins, espe-
cially as nets are reported to be up to 25 km in length
(CCAMLR 2015b). This means that penguin bycatch
data are not available, and it is difficult to make esti-
mates for the entire CCAMLR area without observa-
tions (CCAMLR 2008).
Overall, current penguin bycatch in legal fisheries
within the Convention area is not believed to be a
major threat to Antarctic and Southern Ocean pen-
guin populations. However, IUU fishing, particularly
when large gillnets are used, is a concern.
DISCUSSION
Some level of bycatch has been recorded for 14 of
the 18 penguin species in 4 different gear types (see
Table 2). Despite the above assessments being reliant
on minimal published data, some broad initial find-
ings are clearly identifiable with respect to species
conservation and fisheries management. The impact
of bycatch is not of equal concern for all species
386

Crawford et al.: Penguin bycatch in global fisheries
within a region, varying according to species’ forag-
ing ecologies and behaviours, fishing effort and the
types of gear that predominate within penguin forag-
ing ranges. Because of this, we focus firstly on gear
for which mitigation measures are urgently needed
and secondly on the species towards which limited
conservation resources should be directed.
Gear types of concern
This review highlights gillnets, and to a lesser
extent trawls, as the fishing gear types that most com-
monly lead to penguin bycatch mortality (Table 1).
While interactions have been recorded with long-
lines, the low number of recorded mortalities (with re -
latively high observer effort, particularly in CCAMLR
fisheries) suggests that bycatch in this gear type is a
relatively rare event for all species of penguin.
Penguin entanglements in gillnets are almost in -
evitable given their pursuit diving foraging behav-
iour and the virtual invisibility of modern monofilament
nylon netting when deployed underwater (Martin &
Crawford 2015). Bycatch mortality in gillnets has
been recorded in Magellanic, Humboldt, Galápagos,
African, northern rockhopper, little, Fiordland and
yellow-eyed penguins.
Penguins are capable of deep dives (90 m by Mag-
ellanic penguins, Walker & Boersma 2003; 130 m by
yellow-eyed penguins, Moore et al. 1995), but as air-
breathers, a significant amount of time is spent at the
surface. Additionally, both Magellanic and Hum-
boldt penguins feed on pelagic prey (Frere et al.
1996, Herling et al. 2005) which is a potential reason
for the higher bycatch rates recorded for these spe-
cies in surface-set driftnets compared to bottom-set
gillnets (Majluf et al. 2002, Cardoso et al. 2011). In
contrast, yellow-eyed penguins are predominantly
benthic foragers (87% of dives; Mattern et al. 2007)
and thus are caught in bottom-set gillnets. Gillnets
pose a significant threat to penguins irrespective of
the depth at which they are deployed, and are there-
fore the fishing gear type of greatest concern.
Trawl interactions have been primarily recorded
for Magellanic penguins, but both gentoo and king
penguins have also been captured. While these inter-
actions are subtly different among species, they pri-
marily occur during hauling. Magellanic penguins
are caught as they attempt to forage on smaller fish
dropping out of the net as it is hauled (González-
Zevallos & Yorio 2006), and some of the gentoo pen-
guins caught in the Falkland Islands were actively
feeding on discards (Roux et al. 2012), suggesting
that discard management measures could mitigate
against such bycatch, at least for these species.
The direct impact of purse seine fisheries on pen-
guins has been subject to almost no research, but the
limited amount of data from Chile suggest that, at
least there (where effort is high), the issue of penguin
(and other seabird) bycatch warrants further investi-
gation (ATF-Chile unpubl. data).
Species of concern
To further identify priorities for action on bycatch,
the species that are presently most at-risk from cap-
ture in fisheries are highlighted. Note that this is
based on present scenarios, with limited available lit-
erature on bycatch; these categorisations may change
with new information and changing fishing effort or
species distributions.
High risk
The 3 species for which there is a substantive
amount of evidence of fisheries bycatch are Hum-
boldt, Magellanic and yellow-eyed penguins. The
distributions and gear types of concern for these spe-
cies are shown in Fig. 3.
Magellanic penguins are the best-researched spe-
cies with regard to bycatch (Gandini et al. 1999,
Yorio & Caille 1999, González-Zevallos & Yorio 2006,
Cardoso et al. 2011, González-Zevallos et al. 2011,
Marinao et al. 2014). This species is listed as Near
Threatened, owing to declining trends in the Argen-
tinean breeding colonies that represent the majority
of the estimated global population (Table 2; BirdLife
International 2017). So while no single fishery is
thought to have population-level impacts on the spe-
cies, this trend highlights the concern of potential
cumulative effects of bycatch in various trawl and
gillnet fisheries from breeding colonies in southern
Chile and central/southern Argentina to non-breed-
ing grounds in Uruguayan and Brazilian waters
(Schlatter et al. 2009, Pütz et al. 2011, Stokes et al.
2014). Further, the evidence from some studies (Car-
doso et al. 2011, Marinao et al. 2014, Gianuca et al.
2017) that bycatch is predominantly comprised of
adults increases the possibility that bycatch impacts
are significant at a population level.
Although fewer data are available on Humboldt
penguin bycatch, the IUCN Red List status of this
species (Vulnerable, owing to an overall reduction in
the number of breeding colonies indicating an ongo-
387

Endang Species Res 34: 373– 396, 2017
ing rapid decline; BirdLife International 2017) and
the high number of artisanal fisheries using gillnets
within the species’ core range in Chile and Peru
(Goya et al. 2011) puts Humboldt penguins at risk. In
light of the apparently heightened risk from driftnets
(versus bottom-set nets; Majluf et al. 2002), priority
should be afforded to collecting data in these fish-
eries operating within at least 35 km of key colonies
(Culik & Luna-Jorquera 1997, Luna-Jorquera &
Culik 1999).
Bycatch mortality is of greatest concern for the
Endangered yellow-eyed penguin (Darby & Dawson
2000, Ellenberg & Mattern 2012, Mattern et al. 2017).
The population is decreasing and is now thought to
number fewer than 400 pairs for the New Zealand
mainland population (only 216 nests found during
2015/16 searches; M. Young pers. comm.). Current by -
catch estimates of 35 birds yr−1 (range 16−60; Richard
& Abraham 2015) are potentially an underestimate
due to the lack of independent observer coverage on
commercial gillnetters operating in important pen-
guin foraging areas. However, given the species’
precarious situation, even low numbers of bycaught
penguins will contribute to the species’ demise. Gill-
net effort peaks in summer, which coincides with the
yellow-eyed penguin breeding season. The loss of 1
parent usually results in breeding failure (Darby &
Dawson 2000), and the surviving bird
will generally skip at least 1 breeding
season following the loss of its partner
(Setiawan et al. 2005), further exacer-
bating the impact of by catch.
Moderate risk
Fiordland, Galápagos and little pen-
guins are considered to be moderately
susceptible to bycatch at present. While
only 3 Fiordland penguins have been
officially recorded as bycatch, recent
evidence suggests that bycatch occurs
more regularly, particularly off Stew-
art Island. Limited observer coverage
means there is considerable un certainty
about potential impacts. Given the for-
aging overlaps with commercial gill-
netters and shared foraging grounds
with yellow-eyed penguins, further
observer effort is required to better
determine the risks for this species.
There are no legal fisheries operat-
ing around the Galápagos Islands im -
pacting Galápagos penguins, but illegal gillnetting is
suspected to occur, and this is of some concern, espe-
cially as the species is listed as Endangered (BirdLife
International 2017).
Little penguins are susceptible to capture in gill-
nets, but with few observer data and a relatively
secure conservation status (Least Concern, BirdLife
International 2017), we have ranked the species as
moderately susceptible, pending further studies.
However, if little penguins are split into separate
New Zealand and Australian species (Grosser et al.
2015), this would increase conservation concern and
therefore the relative priority of the bycatch issue.
Low risk
Given the high level of observer coverage in
CCAMLR fisheries and low levels of bycatch re -
corded, the Antarctic species (king, gentoo, chinstrap,
emperor and Adelie penguins) can be considered of
lowest concern with regard to bycatch; indeed, the
latter 2 species have never been recorded as bycatch.
Also of lower concern are Snares and royal penguins,
for which there are no records of incidental captures
in fisheries, most likely linked to the remote locations
of colonies and restricted ranges (but note that
388
Fig. 3. At-sea distribution of the 3 penguin species for which bycatch is cur-
rently of most concern, and the fishing gear types that birds interact with most.
These ranges represent priority sites for conservation action on penguin by-
catch. For yellow-eyed penguin, dark red areas denote foraging range in a
‘normal’ breeding season, with the lighter shade representing expanded
range for breeding birds when food supply is poorer. The pink shading de-
notes the non-breeding foraging range, limited by the 150 m depth contour.
Blue shading: breeding and non-breeding range for Humboldt penguins; or-
ange shading: breeding and non-breeding range for Magellanic penguins.
Source: BirdLife International and Eudyptes Ecoconsulting

Crawford et al.: Penguin bycatch in global fisheries
Snares penguins do forage in areas with active gill-
net fisheries; Mattern et al. 2009, Mattern 2013b).
While erect-crested, macaroni and northern and
southern rockhopper penguins have been recorded
as bycatch, similarly to Snares and royal penguins,
the remote, restricted locations of colonies and lim-
ited presence of these species in areas of high coastal
fishing effort suggests that these species are cur-
rently of least concern. With no evidence of trawl
interactions and gillnet exclusions around colonies
(South Africa) or across the coast (Namibia), the risk
of bycatch for African penguins is presently low,
though this would change if gillnet restrictions were
lifted, given previous bycatch events in this gear.
Importance of bycatch compared to other threats
for the most susceptible species
Poor observer coverage in some areas, including
some commercial inshore fisheries operating in key
penguin foraging areas, impedes the assessment of
the relative importance of fisheries-related mortality
against other threats. However, given limited avail-
able conservation resources, some comparison gives
context and aids prioritisation. Climate change is
likely to affect all of these species, albeit in different
ways (García-Borboroglu & Boersma 2013, Trathan et
al. 2015, Mattern et al. 2017). In order to improve
resilience of penguin populations in the face of cli-
mate change, other threats need to be substantially
reduced, and incidental mortality in fishing gear is
one threat that can be addressed immediately.
The greatest threats to Magellanic penguins are at
sea (García-Borboroglu & Boersma 2013), and of
these, bycatch is perhaps the most significant. No
industrial fisheries are currently in competition with
Magellanic penguins for the same prey species (con-
trary to Humboldt and African penguins; García-Bor-
boroglu & Boersma 2013). Oil spills have been of con-
cern (García-Borboroglu et al. 2008), and although
they have decreased in frequency in recent years,
they should still be considered with fisheries bycatch
in terms of potential population level effects.
The greatest threat to Humboldt penguins is be -
lieved to be competition with fisheries for Peruvian
389
South America Southern
Africa Australasia Antarctica
Species
Population size
estimate
IUCN Status
Argentina
Falklands/Islas
Malvinas
Uruguay
Brazil
Chile
Peru
Ecuador
South Africa
New Zealand
Australia
Antarctica &
Southern
Ocean Islands
Magellanic 1.1–1.6 million pairs NT T; G T T; PS G G; PS
Galápagos 1200 ind. EN G
Humboldt 32000 ind. VU GG
African 50000 ind. EN G
Little 469760 ind. LC GG
Southern
rockhopper 2 500000 ind. VU LL LL
Northern
rockhopper 240300 pairs EN G
Macaroni 6.3 million pairs VU LL
Fiordland 5500–7000 ind. VU G
Erect-crested 150 000 ind. EN LL
Yellow-eyed 1700 breeding pairs EN G
Chinstrap 4 million pairs LC LL
Gentoo 774 000 ind. LC T; LL LL
King 1.6 million pairs LC T LL; T
Table 2. Recorded penguin bycatch in fisheries. IUCN status and population size estimate from BirdLife International (2017).
EN: Endangered, VU: Vulnerable, NT: Near Threatened, LC: Least Concern. Gear types are G: gillnet, LL: longline, PS: purse
seine, T: trawl. Grey boxes indicate that country is not part of that species’ range

Endang Species Res 34: 373– 396, 2017
anchoveta Engraulis ringens (Herling et al. 2005),
particularly in combination with impacts on prey
resulting from more frequent El Niño-Southern Oscil-
lation events (Culik et al. 2000). Humboldt penguins
are vulnerable to unregulated tourism, but its impacts
have not been quantified at a population level (Ellen-
berg et al. 2006). Furthermore, some Humboldt pen-
guins are taken for illegal consumption and bait both
in Chile and Peru (P. McGill pers. comm.). Alfaro-
Shigueto et al. (2010) noted that mortality of Hum-
boldt penguins could be exacerbated by the reten-
tion of incidentally captured penguins for human
consumption, and Trathan et al. (2015) mentioned
egg collection as a threat. As this review has docu-
mented, Humboldt penguins are bycaught in gillnets
(Majluf et al. 2002, Pro Delphinus unpubl. data). As
many vessels use this gear within the Humboldt pen-
guin foraging range (Goya et al. 2011), the region is
clearly a priority for further research and mitigation,
in association with ecosystem-based management of
the anchoveta fishery and ensuring that tourism is
carefully managed at colonies.
The situation is complex for yellow-eyed penguins;
the population is small and decreasing, and is facing a
wide spectrum of threats (Seddon et al. 2013, Mattern
et al. 2017), including sensitivity to human disturbance
(Ellenberg et al. 2007, 2013), vulnerability to disease
outbreaks and marine pollution, the loss of mature
coastal forest as breeding habitat, and predation by
introduced mammals (Seddon et al. 2013). Further-
more, recent research shows that as sedentary benthic
foragers, yellow-eyed penguins are particularly vul-
nerable to climate change (Mattern et al. 2017).
Whereas terrestrial threats are reasonably well
understood and managed, threats at sea require urgent
attention. There are marked natural inter-annual
variations in food availability affecting reproductive
success and survival, with 33% of population vari-
ability explained by ocean warming alone (Mattern
et al. 2017). The impact of climate change is expected
to intensify, making populations less resilient to non-
climate related impacts. In addition to bycatch in gill-
nets, there is concern about the potential indirect
effects of habitat degradation from commercial dredg-
ing (Browne et al. 2011) and bottom trawl fisheries
(Mattern et al. 2013) impacting on prey species, and
the indirect competition from overfishing of spawn-
ing stocks (Ellenberg & Mattern 2012). Similar to by -
catch, these other fisheries-related threats are poorly
quantified. Of the at-sea threats, reducing bycatch
mortality of yellow-eyed penguins is the most feasi-
ble action to improve the resilience of this Endan-
gered species.
RECOMMENDATIONS
The following recommendations are offered to re -
duce penguin bycatch and to provide the critical data
required for evidence-based management.
Species-specific monitoring and bycatch estimates
Given the limited available data on penguin by -
catch, and the potential threat posed to the 3 priority
species listed above, there is a strong case for improved
bycatch monitoring in key fisheries. Independent
monitoring of fisheries operating in penguin foraging
areas is essential to quantify actual bycatch mortality.
Although limited resources may reduce the ability to
monitor fleets via costly observer programmes, by -
catch of other taxa (i.e. cetaceans) has been success-
fully, and more cheaply, monitored by video camera
(Kindt-Larsen et al. 2012). The use of electronic mon-
itoring in surface longline fisheries in Australia has
significantly modified fisher behaviour with respect
to logbook reporting, including seabird bycatch (Lar-
combe et al. 2016). The small size of many gillnet
vessels means there are difficulties in placing physi-
cal observers on board, further strengthening the
case for higher levels of electronic monitoring.
Monitoring priorities for the 3 most at-risk species
are listed in Box 1 below.
In addition, significant data gaps exist for species
for which single instances of bycatch have been re -
corded, meriting further investigation. Specifically,
the extent of gillnet use in South Africa and Namibia
and the risk to African penguins in these fisheries
must be addressed. Monitoring of gillnet fisheries is
required in Bass Strait, Australia, in light of the num-
ber of little penguin captures. Additionally, the con-
servation status of the Galápagos penguin warrants
closer monitoring of illegal gillnet fisheries.
Technical mitigation measures
Effective technical mitigation measures are yet to
be fully developed to reduce sub-surface seabird
interactions with gillnet and trawl fisheries. Some
measures have shown promise, including thicker,
white meshes in the upper sections of driftnets and
the use of acoustic ‘pingers’ to deter auks (Melvin et
al. 1999). Recent research has proposed the deploy-
ment of black and white panels to alert birds to gill-
nets (Martin & Crawford 2015), and tests of LED
lights clipped along the headline of gillnets have
390

Crawford et al.: Penguin bycatch in global fisheries
reduced turtle (Wang et al. 2013, Ortiz et al. 2016)
and seabird (Mangel et al. unpubl. data) bycatch.
Adjusting the twine colour of surface gillnets (in less
turbid waters) may also have some utility (Hanam-
seth et al. in press). Further research into potential
technical measures should prioritise fisheries with
known penguin interactions.
Site identification and spatial/temporal closures
In the absence of established mitigation measures,
and given the elevated conservation concern for many
penguin species, identification of the most important
at-sea sites for penguins (e.g. via tracking where pos-
sible or using foraging radii around colonies) will help
to identify where penguin foraging areas overlap
with fisheries and inform MPAs, with appropriate spa-
tial or temporal fisheries closures. The higher rates of
Magellanic penguin bycatch observed closer to shore
for colonies in Argentina (Yorio et al. 2010, González-
Zevallos et al. 2011, Marinao et al. 2014) emphasises
the potential for spatial approaches. Seasonal or finer-
scale temporal closures in particular may have poten-
tial for Magellanic (González-Zevallos et al. 2011)
and Humboldt penguins (Majluf et al. 2002), and
could take advantage of the existing system of MPAs
in the region.
The temporal/spatial fisheries management and
the creation of MPAs is particularly urgent for yel-
low-eyed penguin, the most threatened species con-
sidered here. Eighteen important sites have already
been identified (Forest & Bird 2015); yet, thus far, no
MPAs have been established in important yellow-
eyed penguin foraging areas.
The majority of the 290 Important Bird and Biodi-
versity Areas identified for penguins to date are ter-
restrial (BirdLife International 2016), making the
identification of at-sea areas particularly important to
inform fisheries and other conservation management
at sea.
Acknowledgements: This review was conceived at a bycatch
workshop at the 8th International Penguin Conference in Bris-
tol, UK, and we are grateful to participants at the workshop
for their suggestions, data and comments. The review greatly
benefited from the insightful comments of P. Dee Boersma,
Luis A. Cabezas, Pablo García-Borboroglu, F. Hernán Vargas,
Dave Houston, Tamara Martínez, Patty McGill, Bruce Mc -
Kinlay, Godfrey Merlen, Jaime Ojeda, Norman Ratcliffe,
Philip J. Seddon, Bryan Wallace, Ralph Vanstreels and Oliver
Yates, as well as 2 reviewers. Thanks to the Galápagos
National Park Directorate, Charles Darwin Foundation and
donors for data on Galápagos penguins. This research did
not receive any specific grant from funding agencies in the
public, commercial or not-for-profit sector.
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Editorial responsibility: Brendan Godley,
University of Exeter, Cornwall Campus, UK
Submitted: February 21, 2017; Accepted: October 7, 2017
Proofs received from author(s): November 22, 2017