Received: 5 October 2022 Revised: 15 February 2023 Accepted: 5 March 2023

DOI: 10.1111/raq.12811

REVIEW

A global review of protected species interactions with

marine aquaculture

Gretchen E. Bath 1 | Carol A. Price 2 | Kenneth L. Riley3 | James A. Morris Jr3

1
CSS, Inc. on contract with NOAA National
Ocean Service, National Centers for Coastal Abstract
Ocean Science, Beaufort, North Carolina, USA
As marine aquaculture expands around the world, there is increased concern for
2
JHT on contract with NOAA National Ocean
Service, National Centers for Coastal Ocean impacts to protected species. Documented cases of marine mammal, sea turtle,
Science, Beaufort, North Carolina, USA seabird, and shark interactions with aquaculture installations do exist but are
3
NOAA National Ocean Service, National
challenging to find. This extensive review summarises the state of knowledge of pro-
Centers for Coastal Ocean Science, Beaufort,
North Carolina, USA tected species interactions with marine fish and shellfish aquaculture installations.
Although seaweed aquaculture was beyond the scope of this review, some of the
Correspondence
Gretchen E. Bath, CSS, Inc. on contract to findings for shellfish and finfish aquaculture may be relevant. The potential impacts
NOAA National Ocean Service, National of farms including habitat exclusion, entanglement, entrapment, collisions, and beha-
Centers for Coastal Ocean Science, 101 Pivers
Island Rd., Beaufort, NC 28516, USA. vioural modifications are the primary risks posed to protected species by marine
Email: [email protected] aquaculture facilities and operations. In addition, indirect effects from habitat impacts
Present address of farm operations may be of concern in some areas as well as the cumulative
Carol A. Price, North Carolina Aquariums and
impacts of multiple small or large farms in the same general vicinity. Decades of farm
North Carolina State University Center for
Marine Sciences and Technology, Morehead innovations and best management practices have been driven by industry, natural
City, North Carolina, USA.
resource managers, conservation organisations, and international conservation agree-
ments. This review is useful for informing industry planning and permitting to
develop aquaculture in the open ocean. This work will help advance the science of
conservation by synthesising the state of knowledge and provide managers and
industry with more insight to protect the most vulnerable species.

KEYWORDS
aquaculture interactions, marine cage, marine mammals, mussel longline, protected species,
sea turtles

1 | I N T RO DU CT I O N a diverse number of feed alternatives6; access to ports; a stable legal

and economic system; skilled labour; and substantial seafood market
The increasing demand for seafood and marine aquaculture techno- demand.1–4,7
logical innovations of the last three decades provide an opportunity to To assess the potential effects aquaculture activities could have on
increase global production of protein-rich, nutritious seafood in the protected species, regulatory authorities rely on the best available sci-
ocean.1–5 A study by the United Nations Food and Agriculture Orga- entific information, which includes but is not limited to peer-reviewed
nization (FAO) identified significant marine aquaculture growth poten- scientific data and publications. Herein, we summarise historical records
tial with high opportunity for farming within exclusive economic of protected species interactions with marine aquaculture operations
zones.1 In addition, increased seafood production can benefit from globally. Many new data are presented and together represent the
vast coastlines that span from polar to tropical climates with suitable most comprehensive review of documented interactions to inform
depths, current speeds, and temperatures; mature gear technologies; resource conservation and risk assessment associated with marine

© 2023 John Wiley & Sons Australia, Ltd. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.

Rev Aquac. 2023;1–34. wileyonlinelibrary.com/journal/raq 1

2 BATH ET AL.

aquaculture development. This information will be valuable as a starting of the same type of interactions should be considered as aquaculture
place for coastal managers conducting environmental review of marine development moves offshore. And while some environmental con-
aquaculture or in their role providing technical advice or consultations cerns are alleviated, the challenge of monitoring farms for protected
on aquaculture activities that might impact endangered and threatened species interactions may be compounded because of the remoteness
species, marine mammals, and important marine habitats. This review of offshore farming.
summarises the current state of knowledge regarding documented
interactions of marine mammals, sea turtles, seabirds, and sharks with
aquaculture farms worldwide. 2 | MARINE AQUACULTURE GEAR
Over the past 30 years, protected species interactions with aquacul-
ture have been documented in a wide variety of literature including 2.1 | Longline mussel aquaculture
government agency documents, non-governmental reports, journal publi-
cations, and books. Our review included scientific papers; government Coastal and offshore mollusc aquaculture production is predominately
reports and databases; industry organisation documents; company mussels (Mytilidae), sea scallops (Pectinidae) and oysters (Crassostrea
sustainability and investor reports; certification program requirements, spp.).5 Globally, the primary cultivation species are mussels, of which
published data, and reports; newspaper articles and press releases; oper- 40% of modern world mussel production occurs in China.5 Other sig-
ational commercial farms; regional fishery management organisations; nificant mussel producers include Canada, Chile, New Zealand, Spain,
stranding and entanglement response organisations; and international France, and Italy. Farmed mussel culture operations employ different
commissions and conventions. Keyword searches were also conducted technologies depending on the location in intertidal, shallow subtidal
using common scientific literature databases including Web of Science, environments, or waters deeper than 20 m. As mussel aquaculture
ProQuest Aquatic Science Fisheries Abstracts, Elsevier Science Direct, expands away from user conflicts and some environmental concerns
JSTOR, and Google Scholar. Early reviewers and colleagues also provided inshore to more exposed offshore, high-energy environments, sub-
relevant publication recommendations. Professional contacts from merged high-tension longline designs are superior (Figure 1).27–32
aquaculture farms supplied unpublished data. The reviewed documents Mussel farms employ both floating and submerged configurations
originated from research conducted globally, cover a range of cultured depending on the physical environment conditions (Figure 1). Coastal
species, include many new and practical farm management approaches, and offshore commercial scale longline mussel farms deploy both spat
and address ecological processes at multiple scales. Less accessible docu- (shellfish larvae) collection and grow-out ropes in multiple backbone
ments or sources may have been missed due to limits on information lines arrayed in parallel rows with individual lines (droppers) vertically
sharing, language barriers, or policy relevance. Noteworthy is that all suspended or a long continuous looped line.33 Single droppers are
countries do not have the same protections for these species, reporting 3- to 10-m long.33,34 Continuous droppers are up to 10-m long and
incidents is therefore not required in all countries; thus, significant data are looped at intervals along the mainline. Collectively, the culture
gaps were expected. rope may be up to 5-km long at a single farm.30,33–36 In an exercise to
There are no conventions for delineating coastal and offshore quantify the cumulative length of lines in a Prince Edward Island
marine aquaculture.8–13 Generally, coastal farms are visible from the embayment, McKindsey et al.37 estimated the total backlines to be
shoreline, moored at shallower depths, typically between 10 and 50 m, over 500 km and socking material and product to be 2250 km within
and may experience significant wave exposure. Coastal farms include a 7 km2 area.
operations found in large embayments like lochs and fjords, and provide Mussel longline configurations are supported by a set of floats
some protection from open ocean conditions. We considered offshore and secured by mooring lines.29,34,35,38,39 The longlines are typically
farms to include aquaculture operations beyond 3 miles from shore. 150–300 m in length with arrays spaced 10–20 m apart and the configu-
Many offshore farm sites are located at water depths greater than ration anchored 5–20 m below the surface by two or four mooring lines
40 m, and are more exposed and subject to ocean swells, strong winds, and suspended by two corner floats.30,32,40 Vertical lines tether mussel
and strong ocean currents with significant wave heights up to 3–4 m. longline arrays to the seafloor using screw anchors, deadweight anchors,
These farms are generally large-scale commercial enterprises requiring or hydraulic expansion anchors.29,34,35 Longlines and anchor lines may be
significant capital investment due to the technology costs. made of durable synthetic line with diameters approximating 36 mm.23,38
Aquaculture farm design and engineering have advanced the Vertical lines connected to anchors and horizontal longline orientations
capability to withstand dynamic offshore environments and increase are maintained in the water column by buoys, which prevent the lines
production capacity.2,3,12,14–17 Offshore, open ocean waters provide from becoming entangled. To alert traversing vessels, radar-reflecting sur-
space for aquaculture expansion, increased protein production, face buoys are attached to the end of each longline. Additional floats
reduced social conflict, and lower exposure to terrestrial sourced pol- added along the longline compensate for their weight and maintain the
lution.2,18–20 The water depth, currents, and ocean circulation provide longline geometry.35 Submerged longlines are deep enough (5–20 m) to
optimal environmental conditions for growing diverse marine species avoid interaction with navigation. Mussel farm infrastructure components
and the potential to reduce some of the negative environmental that pose potential risk for entanglement and injury include anchor
impacts of coastal fish farming.2,9,21–25 In this review, many of the ani- lines, horizontal backbone longlines, vertically suspended and looped
mal encounter reports took place in traditional coastal farms. The risk grow lines, suspended nets, and surface buoy marker lines.26 The slack
BATH ET AL. 3

F I G U R E 1 Representative schematics of surface (a) and submerged (b) offshore longline systems used for suspension culture of mussels,
pearl oysters, and scallops. View (a) shows a single looped grow rope configuration. View (b) shows individual dropper lines suspended from the
backbone.26

spat-collecting lines, grow-out lines, and surface marker buoy lines have requirements. Cages can be lowered and raised in the water column
specifically been implicated in documented entanglement cases.36,41–43 by mechanical systems or air compressors.
A traditional open cage comprises a cylindrical net with bottom
weights to spread the bag, jumping net fixed above the net bag to pre-
2.2 | Finfish aquaculture vent fish escapes, cage collar for net bag spread and buoyancy to keep
the bag in the correct water column position, and a mooring sys-
Technological innovation continues to expand opportunities for tem.11,17 Finfish cages and antipredator netting are made of rigid, syn-
marine finfish aquaculture farms to move further offshore with rigid thetic materials engineered to tolerate ocean conditions for several
or flexible frames capable of floating or being submerged below the years.11,13,17,48 Net used in cage aquaculture are most commonly
9,11,13,44–46
surface. Floating rigid frames (Figure 2a) are made of made with nylon. Rigid polyethylene terephthalate (PET), copper, and
large rubber and high-density polyethylene collar cages.9,11,46–48 stainless steel mesh are also used. Compared with nylon, these rela-
Square steel cage systems (Figure 2b) provide a rigid platform and tively inflexible materials maintain cage structure better in heavy cur-
infrastructure from which holding pens (nets) can be aligned in any rents.11,46,49 Weights or heavy bottom ring/sinker tubes at the
47
configuration and are secured by one mooring line each. Submers- bottom of the net bag help maintain their shape45 but also increase
ible cage systems, such as the SeaStation (Figure 2c) and futuristic the dynamic forces from waves (stretch and slack) acting on the net
geodesic designs (Figure 2d) are engineered to withstand the harsh, bag.11,13,17,50,51 To prevent bowing or folding in currents, antipredator
16,17,31
high-energy conditions of the open ocean. Pen diameters can nets are stretched taut and are deployed with space between them
range from around less than 10 m up to 100 m (up to 200 m for tuna and the fish containment cages. Tensioned deployment reduces
ranching) depending on the production goals of the farm and the site entanglement risk and prevents predators from biting or pushing into
4 BATH ET AL.

F I G U R E 2 Examples of marine cages for finfish production. (a) Floating surface cage system, (b) square steel cage system, (c) submersible
cage system, and (d) the submersible geodesic cage.16,17,31,47

nets to access fish. Depending on the target species for exclusion, pose potential entanglement risk, resulting in injury, stress, or death
antipredator mesh sizes range from 3.8 to 20.0 cm (bar length).11,26,48 to marine mammals, sea turtles, birds, and sharks.25,26,48,53–59 Finfish
Offshore finfish aquaculture infrastructures employ anchoring operations can attract some piscivorous animals because wild fish
systems similar to those found in mussel farms.29 Ropes are used for aggregate to the structural habitat provided by cages or are attracted
both the mooring lines and grid system lines and are most commonly to excess feed loss from the cages.25,55,60,61
made of polysteel with a tensile strength more than 25% higher than
polypropylene. Polyester or nylon ropes are also used; however, they
stretch when placed under load, which can compromise the mooring 3 | PROTECTED SPECIES AND MARINE
11
grid. Cages are tethered in arrays by bridling systems on a farm site AQUACULTURE
(Figure 3) and often secured by orthogonal moorings. In many cases,
double anchors and more durable lines require sufficient flexibility to 3.1 | Marine mammals
ensure that wind and water forces on net structures do not fully
stretch the lines and generate a full load.11,14,46,52 For some cages Marine aquaculture and marine mammal interactions have occurred
(typically submersibles), single point mooring systems are designed throughout the world.26,55,58 Events are documented in Australia and
with each cage anchored individually with a single line. The cage can Tasmania,60 New Zealand,36,43 United States,62 Canada,63 Argentina,64
rotate in an arc defined by the length of the mooring line. In deep Chile,57,60,65,66 Iceland,67 Scotland,68 Norway,69 Italy,70 Turkey,71 and
water, a single mooring and anchor line may be attached to several South Korea.72 Pinnipeds (seals, sea lions, and walruses) and cetaceans
aquaculture cages. Cage locations and farm boundaries are marked (porpoises, dolphins, and whales) are the groups most commonly docu-
with navigational surface buoys11 that could pose a risk for primary or mented to have direct physical interactions because they are more
secondary animal entanglements if the lines are slack. broadly distributed and must surface to breathe. The primary risks
High-performance netting materials are often used in offshore posed to marine mammals by aquaculture facilities and operations are
aquaculture environments to contain fish, exclude predators, and pre- habitat exclusion, entanglement, and behavioural alterations (attraction,
vent predation. Some netting materials and their related hardware avoidance, or food preference).26,36,43,58,73–75
BATH ET AL. 5

F I G U R E 3 Various systems for mooring ocean cages: single point moorings for individual cages, multi-cage grid systems commonly used for
nearshore cages and platform farms, tension leg moorings that allow the cage to rotate in an arc typically used for submerged cages. Source: Lekang.17

3.1.1 | Habitat modification, exclusion, or predators and alternatively, farm structures may aggregate prey and
competition for space provide novel foraging opportunities.84

Marine mammals can be excluded from their habitats depending on the Mussel farms
size and concentration of farms, farm operations (including vessel traf- Several studies in Admiralty Bay, New Zealand, have focused on
fic), and behaviour of a particular marine mammal spe- potential dolphin habitat exclusion in nearshore waters where mussel
cies.43,53,58,60,65,76–78 While some marine mammals may not be farms are located. During 5 years of observations in a bay with
spatially excluded from farm areas, limited mobility in the vicinity may numerous mussel farms, dusky dolphin (Lagenorhynchus obscurus, Gray
result in individuals being forced into suboptimal foraging habitat. Spe- 1828) groups entered farms (located <200 m from shore) in only eight
cies may alter their behaviour, and be deterred from traversing or feed- of 621 observations.85 Compared with unfarmed areas in the Bay, the
ing if aquaculture structures present a navigation obstacle. Animals may dolphins avoided farm areas and were able to navigate through the
not only avoid the farm, but also the broader area; thus, minimising spa- lanes between the mussel lines. Coordinated dolphin feeding behav-
tial overlap with home ranges, foraging habitats, critical breeding areas, iour appeared limited by the presence of the farms.85,86 Duprey84
and migration routes is prudent for farm site selection. In some cases, reported similar findings in the same area; only two of the 332 groups
farm structures may not have a major impact as sited; however, as mul- of dusky dolphins observed were inside a mussel farm and of the nine
tiple farms are constructed within an area, cumulative impacts over groups of bottlenose dolphins (Tursiops truncatus, Montagu 1821)
time to individuals and populations are possible.79,80 These anthropo- seen, none entered farms. A study of dusky dolphin behaviour simi-
genic landscapes can present both risks and benefits for both prey and larly found the animals increased foraging behaviour adjacent to the
predators.81–83 Prey may use structures for safe harbour to escape farms but did not enter the farms.87 Thus, mussel farms may act or be
6 BATH ET AL.

perceived by dolphins as three-dimensional (3D) obstructions that
pez92 investigated the influence of the shellfish aquaculture indus-
Lo
impede navigation and foraging capabilities at and below the surface. try on the specific foraging behaviours of bottlenose dolphins. The
43
Clement suggested there is a low habitat exclusion risk to mussel rafts provided a physical structure that attracted forage fish
marine mammals at both offshore mussel spat collection sites and species, in comparison to adjacent areas to the farms. Pelagic and
grow-out operations in New Zealand, which is consistent with earlier demersal fish species aggregated around the rafts for shelter and to
reports on environmental impacts of the industry.42,60,88 Clement43 feed directly on the line-associated organism communities. Because
cautioned as long as mussel farming expansion does not overlap with their prey was concentrated in these farms, the dolphins were able to
breeding, migrating, and feeding habitats of protected species, few forage more efficiently by shortening their dive times which increased
negative interactions are expected. Although pinnipeds have been oxygen intake and gave them the ability to regulate their mode of
attracted to nearshore mussel farms because they occasionally con- swimming speed.
sume benthic organisms typically associated with mussel farms, includ-
ing crabs and fish,89 they do not commonly feed on shellfish and may Marine finfish farms
be less likely to visit offshore mussel farms.55,62 Kemper et al.60 evalu- Marine mammals can be attracted to the caged fish as well as the
ated known negative interactions of marine mammals with aquaculture presence of wild fish that congregate around fish farms.25 Off the
in the southern hemisphere, and found most occur at finfish farms and coast of Italy, bottlenose dolphins were observed feeding on wild fish
involve pinnipeds attracted to the gear seeking food. in the vicinity of marine fish farms, but did not target the caged fish.94
In southern Chile, both Peale's dolphins (Lagenorhynchus australis Bottlenose dolphins are regularly observed foraging near fish farm
Peale, 1848) and Chilean dolphins (Cephalorhynchus eutropia Gray, cages in the northern Mediterranean.70,73,95–99 Bottlenose dolphins
1846) observed in extensively farmed areas (shellfish and finfish) modified their social structure and altered hunting tactics in response
avoided direct interaction with farms.57,60,65,66 While Peale's dolphins to increased prey densities around fish farms.70,94 In a 5-year study of
were never observed closer than 100 m to mussel farms, Chilean dol- gilthead sea bream (Sparus auratus Linnaeus, 1758), European sea bass
phins were observed feeding on schooling fish adjacent to farms and (Dicentrarchus labrax Linnaeus, 1758), and meagre (Argyrosomus regius
in open spaces between dense sets of growth lines. Seven Chilean
pez73 observed known
Asso, 1801) cages off the coast of Italy, Díaz Lo
dolphins crossed under shellfish lines and floats; however, the clear- individual dolphins exhibit habitat use patterns and farm fidelity. Dol-
ance between the lines and seafloor was unknown. In mussel farm phin occurrence near the farm varied with time of day, season, and
areas, Ribeiro et al.57 reported Chilean dolphins were present with year. Dolphins near farms typically foraged on wild fish but also fed
less than 30% coverage of mussel farms but notably absent in areas on discarded or escaped farmed fish during harvesting operations.
with greater than 60% coverage. In this region, habitat exclusion due Ninety-nine observation months over 9 years at these cages, Díaz
to high density of aquaculture was considered a concern because it
pez100 witnessed bottlenose dolphins habituating to fish harvesting
Lo
restricted essential habitat access. operations where discarded fish were easy prey. Piroddi et al.97 sug-
5
In Europe, shellfish production is second to Asia and little gest bottlenose dolphin abundance increased around fish farms in
research has addressed impacts to protected species.90 In Bantry Bay Greece because the farms facilitated prey capture. Bonizzoni et al.99
89
on the southwest coast of Ireland, Roycroft et al. assessed impacts observed more bottlenose dolphins in areas within 5 km of fish farms
of mussel culture on common seals (Phoca vitulina Linnaeus, 1758). and less at areas more than 20 km from farms. Dolphins did not
The mussel farms used 15 m vertical grow lines suspended from float- appear to avoid farm structures or noise from farm activities and were
ing longlines in nearshore, sheltered, deep water (up to 20 m). Seal thought to be foraging, often within 10 m or less of the fish cages.
abundance was the same at sites with and without mussel farms, and Interviewed farm employees revealed dolphins were not considered a
no negative interactions were reported. Along the northwestern coast threat and acoustic deterrent devices are not used in the area.
of Spain, mussel farms influence habitat uses and foraging behaviour Farmers in Italy, Spain, Malta, Greece, and Israel claim the animals
of bottlenose dolphins.91,92 The shellfish culture method may impact negatively impact their businesses because of depredation on cultured
animals differently due to the gear configurations and materials used. fish as well as inducing stress on those fish, which heightens the
In contrast to longline systems, rafts support mussels grown on ropes importance of understanding the interactions and developing consis-
tied to rectangular floating platforms. Each individual raft has a maxi- tent mitigation measures.73
mum of 500 ropes (no longer than 12 m) and covers an area of up to In the Bay of Fundy, eastern Canada, feeding harbour porpoises

pez and
500 m2.93 Over 2 years of consecutive fieldwork, Díaz Lo (Phocoena phocoena Linnaeus, 1758) were not displaced by an Atlantic
Methion91 observed increased numbers of bottlenose dolphins at salmon farm, except during short periods of feed delivery or cage clean-
mussel farm locations and in waters surrounding aquaculture zones. ing.101 In the same region, Jacobs and Terhune102 observed harbour
They postulated this positive relationship was due to the available seals (Phoca vitulina Linnaeus, 1758) were not attracted to areas with
high-quality prey from large aggregations of fish around mussel rafts. salmon farms. Cetacean species observed in coastal waters of Scotland
Their observations were different than previous studies that observed include harbour porpoises, minke whales (Balaenoptera acutorostrata
other coastal cetacean species avoiding mussel longline aquaculture Lacépède, 1804), killer whales (Orcinus orca Linnaeus, 1758), and bottle-
zones, which resulted in habitat loss and the possibility of negative nose dolphins.68,103–106 There are no reports of these species interact-
population impacts.55,57,76 Building on this study, Methion and Díaz ing with aquaculture gear; however, harbour seals and grey seals
BATH ET AL. 7

(Halichoerus grypus Fabricius, 1791) are considered primary predatory to additional anthropogenic stressors including gillnet fisheries, boat
species of finfish aquaculture sites.107 traffic, and tourism.111 Minke whales (B. acutorostrata) have been seen
60
Kemper et al. evaluated known negative interactions of marine interacting with finfish farms in Chile, although no specific information
mammals with aquaculture in Australia, Tasmania, New Zealand, and was documented.60,113
Chile and found that most occur at finfish farms and involve pinnipeds Based on an ecosystem modelling approach in the Ionian Sea,
attracted to the gear seeking food. Sepúlveda et al.108 used stable iso- increased productivity from fish farm nutrients has contributed positively
topes to derive dietary data from foraging South American sea lions to bottlenose dolphin populations in the region.97 As an explanatory vari-
(Otaria flavescens Shaw, 1800) around salmon farms in southern Chile. able, the number of fish farms was the main factor used to reconstruct
They integrated sea lion movement patterns, based on satellite telem- the observed trends in dolphin biomass and distribution from 1997 to
etry, with dietary data to characterise the impacts of an abundant and 2008. Rapid transfer of nutrients through the food web in oligotrophic
predictable source of non-native prey on their foraging ecology. There waters has been shown to increase commercial fish biomass and fish
was large variability in individual sea lion spatial ranges and their farms are known to act as attracting devices for forage fish.26
degree of overlap with salmon farms. Based on isotopic analyses,
farmed salmon were one of the most important prey in the study area;
however, the authors explained that these data were potentially con- 3.1.2 | Entanglement
founded by the possibility of sea lions consuming widely-distributed
feral salmon in the area with the same isotopic ratios. In addition, the Physical interactions between marine mammals and offshore aquacul-
degree of spatial overlap did not correlate with the relative contribu- ture farms increase the risk of entanglement in structures such as anti-
tion of salmon in their diets. Their results suggest that even if an indi- predator nets and mooring lines.26,41,43,114 The potential for marine
vidual animal forages around salmon farms, it does not necessarily mammals to become entangled and drown is a predominant
prey on the cultured fish. In a recent review of pinniped and salmon concern,55 especially given the frequency of entanglements in com-
farm interactions over 50 years of industry expansion, Heredia-Azuaje mercial fishing gear115–118 and marine debris.111,119 Entangled animals
109
et al. ascertained although the primary threat to pinnipeds is have lower reproductive success, which results in population level
intended and unintended killing, breeding or foraging habitat alter- effects, especially for small populations.120 In addition, injuries from
ations may also induce behavioural or social change in impacted areas. entanglement can reduce movement, impede feeding ability, cause
Chilean dolphins are endemic to Chile and are listed as near internal injuries from struggling, constrict blood flow, sever append-
110
threatened on the IUCN Red List of Threatened Species. According ages, and cause infections.121 Animals burdened by dragging gear may
57
to Ribeiro et al., Chilean dolphin movement patterns and habitat use be disconnected from social interactions and communications. While
did not seem influenced by or directly altered by the presence of spatial overlap of farms and habitats increases the risk of interacting,
salmon farms. Heinrich and Reeves110 also found no significant asso- marine mammals can be attracted to the structures that house poten-
ciation identified between areas of intense habitat use and distance to tial prey or seek out aggregating wild fish near the farm sites, which
salmon farms by Chilean dolphins. However, in the fjords of southern increases the opportunity for entanglement.55,60,98,122,123
Chile, salmon farm structures are commonly close to the shore, and Marine mammal interactions with marine aquaculture gear may
Chilean dolphins have been observed avoiding farm cages.57,66,111 depend on several factors. Young, naïve animals are typically more at
While active avoidance minimises direct interactions with the aqua- risk of entanglement, compared with adults because of their inquisi-
culture gear, Chilean dolphin habitat selection is limited to shallow tive nature and inexperience.124,125 Whales and dolphins that use
waters and thus water depth is a principal environmental feature that echolocation to navigate their environment and feed (Odontocetes)
shapes their distribution.111 This close association with shallower may be better able to detect and avoid 3D farm structures compared
water depth may, therefore, put them at heightened risk of negative with species that do not echolocate (Mysticetes); however, if they are
impacts from aquaculture operations sited at inshore locations. This attracted to the abundance of prey in and surrounding cages they can
may also be true for other Cephalorhynchus species including Hector's still become entangled. Larger, less agile species with flippers and fins
dolphin (C. hectori P.-J. van Bénéden, 1881), Commerson's dolphin that extend out from the body42 and species with feeding strategies
(C. commersonii Lacépède, 1804), Heaviside's dolphin (C. heavisidii that involve engulfing huge volumes of water (e.g., baleen whales
Gray, 1828), and Peale's dolphin, which show the same general habitat including right, minke, and humpback whales) are considered more
requirements.66,111 Heinrich et al.66 further refined the broadly sym- susceptible to entanglement in ropes and lines.67 Species or individ-
patric Chilean and Peale's dolphin coastal water habitat requirements uals that roll when encountering entangling gear may be more likely
using species distribution models (SDM). Chilean dolphins preferred to become severely wrapped.126 Marine mammal perception of struc-
shallow (<30 m depth) turbid waters within 500 m of shore and river tures in the ocean and use of visual, auditory, or other sensory cues to
mouths near shellfish farms. Peale's dolphins were also observed in elicit risk-averse behaviour is not well understood.124,125 Not all coun-
shallow waters but occurred over a broad range of conditions along tries mandate marine mammal protection and aquaculture farms may
open and exposed coastlines. With the expansion of aquaculture in not be required to report interactions. Most of the global marine
these shallow, coastal habitats,112 populations of these near-threatened aquaculture occurs in countries with no reporting. Thus, entanglement
species110 become fragmented and isolated, and thus more vulnerable data are relatively sparse and rarely quantitative for both shellfish and
8 BATH ET AL.

TABLE 1 Documented cetacean entanglements or entrapments at mussel farms.

Species
Location common name Scientific name Number Year Gear type Outcome Citation(s)
a
Argentina Southern right whale Eubalaena australis 1 2011 Suspected mussel Unknown 64
(juvenile) farm spat line
Western Australia Humpback Megaptera 1 2005 Mussel farm spat line Released 43,67,127–129
whale (calf ) novaeangliae
Iceland Harbour porpoise Phocoena phocoena 1 1998 Mussel farm spat line Fatal 67
Iceland Humpback whale Megaptera 1 2010 Mussel farm spat line Fatal 67
(juvenile) novaeangliae
Iceland Minke whale Balaenoptera 1 2012 Mussel farm head Released itselfa 67
acutorostrata rope
New Zealand Bryde's whale Balaenoptera edeni 1 1996 Mussel farm spat line Fatal 36,43,67,88
brydei
New Zealand Bryde's whale Balaenoptera edeni 1 Prior to 2003 Mussel farm spat line Fatal 36
brydei
New Zealand Humpback whale Megaptera 1 2011 Mussel farm line as a Releasedb
(calf) novaeangliae result of primary
craypot
entanglement
rescue operation
South Korea North Pacific Eubalaena japonica 1 2015 Mussel farm spat line Released itself 67,72,130
right whale
South Africa Bryde's whale Balaenoptera edeni 1 2019 Mussel farm spat line Released itself 131
brydei
a
Unconfirmed event.
b
https://www.newshub.co.nz/environmentsci/mussel-farmer-frees-trapped-humpback-whale-2011070917.

finfish aquaculture farms in many countries and the global extent of (Table 1). In 2005, a humpback whale calf in Western Australia was
the problems are poorly known. cut free from a mussel spat line after catching it in its mouth and roll-
ing43,67,127–129 (Table 1). In a report on southern right whale (Euba-
Mussel farms laena australis Desmoulins, 1822) entanglements in Argentina from
It is unclear whether entanglement occurs because mammals are 2001 to 2011, there is one unconfirmed case of a juvenile right whale
attracted to or unaware of shellfish-farming gear.58 Lloyd36 reported that may have involved mussel spat collection lines64 (Table 1).
two fatalities of Bryde's whales (Balaenoptera edeni brydei Anderson, The whale was sighted during a whale-watching cruise and was not
1879) entangled in mussel spat collection lines in New Zealand re-sighted during the search effort and the fate is unknown. Two
(Table 1). In 1996, a Bryde's whale was found dead with the spat line fatal marine mammal entanglements in mussel single dropper spat col-
lodged tightly through the base of the animal's mouth indicating that lection lines were reported in Iceland67 (Table 1). In 1998, a harbour
36,43,67,88
the entanglement occurred with a high level of force. There porpoise was found entangled, and in 2010 a juvenile humpback
are no additional details about the second Bryde's whale fatality men- whale entanglement was reported. In 2012, a mussel farmer in Iceland
tioned in Lloyd.36 Bryde's whales are designated Threatened- suspected a minke whale was entangled in headropes and droppers
Nationally Critical in New Zealand132 and since these two reported but it freed itself without gear attached. Although the farmer had not
incidents, no other entanglements have been documented. In 2011, actually seen the whale, he assumed it happened because the gear
an entanglement occurred because of a rescue operation when a was in disarray and slime-coated, similar to what is left on his gillnets
humpback whale (Megaptera novaeangliae Borowski, 1781) calf was after a whale had been caught.67 In 2015, a young male North Pacific
dragging craypot gear tied around its fluke.133 The New Zealand right whale (Eubalaena japonica Lacépède, 1818) was entangled by a
Department of Conservation workers tied buoys to the whale as the mussel spat line in South Korea (Table 1). Divers cut the ropes but had
sun was setting so they could follow the whale and locate it in the to cease operations due to low visibility at night. When they returned
morning when they had better visibility. The next morning a mussel the next morning, the whale was nowhere to be found and assumed
farmer discovered the whale entangled in mussel ropes by the buoy released.67,72,130
lines. He and his crew cut the lines off the whale and freed it without Similar to mussel longline construction, pearl oysters (Pinctada
the offered assistance of trained responders, and unfortunately, this maxima) are held in net panels just below the surface and attached to
was achieved by winching the tail of the whale out of the water133 surface longlines anchored at each end (Figure 1). This allows the
BATH ET AL. 9

TABLE 2 Documented cetacean entanglements at oyster pearl farms.

Species
Location common name Scientific name Number Year Gear type Outcome Citation(s)
Western Australia Humpback whale Megaptera 1 1998 Pearl farm rope Released 128,129
novaeangliae
Western Australia Humpback whale Megaptera 1 2004 Pearl farm rope Released 128,129
novaeangliae
Western Australia Humpback whale Megaptera 1 2008 Pearl farm rope Released 128,129
novaeangliae
Philippines Short-finned pilot Globicephala Not quantified Pearl farm rope 135–137
whale macrorhynchus
Philippines Pantropical spotted Stenella attenuata Not quantified Pearl farm rope 135,136
dolphin

panels to move with the tide and the oysters to feed in as close as dolphins have drowned in antipredator nets that were not enclosed at
possible to their natural state.134 Pearl oyster farm ropes in Western the bottom; trapping the dolphins between the main cages and the
Australia have been implicated in three humpback whale entangle- antipredator nets60,154 (Table 3). Before 2000, 36 Australian fur seals
128,129
ments in 1998, 2004, and 2008; all animals were released alive (Arctocephalus pusillus Wood Jones, 1925), one New Zealand fur seal
(Table 2). In the Philippines, short-finned pilot whales (Globicephala (Arctocephalus forsteri Lesson, 1828), two leopard seals (Hydrurga leptonyx
macrorhynchus Gray, 1846) and pantropical spotted dolphins (Stenella de Blainville, 1820), and one southern elephant seal (Mirounga leonina Lin-
attenuata Gray, 1846) entangled in pearl farm lines135,136 (Table 2) naeus, 1758) died in Atlantic salmon aquaculture farms60 (Table 4). Tassal
although no details about specific events or the cumulative number of Group is the largest Atlantic salmon producer in Tasmania. Since achiev-
animals impacted have been documented. ing the Aquaculture Stewardship Council certification in 2014 across its
Atlantic salmon farm operations in Tasmania, it has publicly released
Offshore finfish farms information about wildlife interactions at all of its sites within 30 days of
From 1990 to 1999, fatal entanglements of 17 short-beaked common occurrence on its sustainability dashboard website.160 These interactions
dolphins (Delphinus delphis Linnaeus, 1758) and 11 Indo-Pacific bottle- are also documented in its annual Sustainability Reports.161 From 2010
nose dolphins (Tursiops aduncus Ehrenberg, 1833) in southern bluefin to 2021, 70 accidental Australian fur seal deaths were recorded in their
tuna (Thunnus maccoyii Castelnau, 1872) ranching operations using salmon net pens (Table 4). From 2014 to 2020, there have been no inter-
large mesh (>15 cm) antipredator nets have been documented in actions with sharks, whales, or dolphins at their farms. Two New Zealand
South Australia53,60 (Table 3). In 1993, a humpback whale broke fur seals became entangled and drowned in finfish antipredator nets in
through the walls of a tuna ranching operation and was trapped in the New Zealand prior to 200743,56 and another two in 2014 (Table 4). In
net pen for 2 days before it was successfully released53,60 (Table 3). In New Zealand, fish farms use antipredator nets to deter pinnipeds; how-
an International Whaling Commission Scientific Committee National ever, these nets have been implicated in small cetacean entanglements.78
138
Progress Report, the fatal entanglement of a bottlenose dolphin in Three dusky dolphins and one Hector's dolphin have been fatally
a yellowtail kingfish (Seriola lalandi Valenciennes, 1833) cage in 2008 entangled.56,78,84,85,148 Four bottlenose dolphin deaths occurred before
was documented with no further information (Table 3). Twenty-one 201343 (Table 3). Huon Aquaculture162 reported 18 seal deaths in 2021
unspecified seal and sea lion interactions were reported at yellowtail on their sustainability dashboard.
kingfish farms and two seal interactions reported in southern bluefin A Bryde's whale measuring 6 m in length was rescued from being
tuna ranching operations from 2014 to 2016. All reported pinniped trapped in a fish farm in Brunei Bay near Pulau Pelompong Brunei-
interactions resulted in the animals releasing themselves or being Maura in May 2000145,146 (Table 3). The whale was injured and had a
155
released alive (Table 4). Over the past 20 years, improvements in the partially severed fluke, not because of the fish cage, but because two
tuna ranching industry with operations and best practices have elimi- fishermen tied a rope around the base to help pull the whale to dee-
nated marine mammal entanglement mortalities.159 Predator nets are per water. The rescuers were able to move the whale to deeper water
no longer used by the tuna industry and additional measures have been and watch it swim away, reportedly struggling to orient itself from the
taken such as switching to pellet feed and reducing waste, better siting damaged tail.136,145,146
159
practices, reduced tuna mortalities and prompt carcass removals. Sea lion mortality in Chile salmon farms is rampant due to shoot-
Atlantic salmon (Salmo salar Linnaeus, 1758) net pens are the most ing, poisoning, clubbing, and some incidental entanglement.60,156,157
abundant aquaculture gear in Tasmania. Before 1991, a southern right In Southern Chile, there is a well-known negative interaction between
whale collided with the side of an Atlantic salmon net pen but was South American sea lions and salmon farms. The high concentration
released, most likely after getting tangled in mooring lines, not preying on of vulnerable prey results in the threat of predatory attacks at the
the caged fish153 (Table 3). Bottlenose and short-beaked common cage which contributes to farmed salmonid escapes.156,163 Pinniped
10 BATH ET AL.

TABLE 3 Documented cetacean entanglements or entrapments at marine finfish farms.

Species
Location common name Scientific name Number Year Gear type Outcome Citation(s)
Australia Bottlenose Tursiops truncatus 1 2008 Kingfish cage Fatal 138
dolphin
South Australia Humpback whale Megaptera novaeangliae 1 1993 Tuna ranching Released 53,60
operation
South Australia Short-beaked Delphinus delphis 17 1990–1999 Tuna ranching Fatal 53,60
common dolphin operation
South Australia Indo-Pacific Tursiops aduncus 11 1990–1999 Tuna ranching Fatal 53,60
bottlenose operation
dolphin
British Harbour porpoise Phocoena phocoena 4 2007 Net pen Fatal 139
Columbia
Canada
British Humpback whale Megaptera novaeangliae 1 2013 Net pen Found dead at 139
Columbia farm
Canada
British Humpback whale Megaptera novaeangliae 2 2016 Net pen Fatal 139
Columbia
Canada
British Humpback whale Megaptera novaeangliae 1 2016 Net pen Released 139
Columbia
Canada
British Humpback whale Megaptera novaeangliae 1 2018 Net pen Releaseda 139
Columbia
Canada
Chile Chilean dolphin Cephalorhynchus eutropia 1 2007 Net pen Fatal 59,140
Chile Humpback whale Megaptera novaeangliae 1 2007 Net pen Fatal 141,142
(calf)
Chile Chilean dolphin Cephalorhynchus eutropia 1 2011 Net pen Fatal 59
Chile Chilean dolphin Cephalorhynchus eutropia 4 2015–2017 Net pen Fatal 59
Chile Humpback whale Megaptera novaeangliae 1 2017 Net pen Released 59
(adult)
Chile Sei whale Balaenoptera borealis 1 2020 Salmon net pen Fatalb
Iceland Minke whale Balaenoptera 1 2005 Net pen Fatal 67,143,144
acutorostrata
Brunei Bryde's whale Balaenoptera edeni brydei 1 2003 Finfish cage Released 136,145,146
Italy Bottlenose Tursiops truncatus 3 2005 Coastal fish farms Fatal 70
dolphin
New Zealand Hector's dolphin Cephalorhychus hectori 1 1987 Net pen Fatal 147
New Zealand Dusky dolphin Lagenorhynchus obscurus 7 1999–2018 Net pen Fatalc 56,78,84,85,
87,147,148
New Zealand Hector's dolphin Cephalorhychus hectori 1 2005 Net pen Fatal 56,147,149
New Zealand Bottlenose Tursiops truncatus 4 Prior to 2013 Net pen Fatal 53
dolphin
Norway Humpback whale Megaptera novaeangliae 1 2009 Net pen Released 144,150
Norway Minke whale Balaenoptera 1 2005 Net pen Fatal 67,143,144
acutorostrata
Norway Minke whale Balaenoptera 1 2014 Net pen Released 151
acutorostrata
Norway Humpback whale Megaptera novaeangliae 1 2015 Net pen Releasedd 151
(calf)
Norway Minke whale Balaenoptera 1 2015 Net pen Fatal 151
acutorostrata
BATH ET AL. 11

TABLE 3 (Continued)

Species
Location common name Scientific name Number Year Gear type Outcome Citation(s)
e
Norway Harbour porpoise Phocoena phocoena 1 2018 Net pen Fatal
Norway Minke whale Balaenoptera 1 2019 Net pen Unknownf 151,152
acutorostrata
Tasmania Southern right Eubalaena australis 1 Prior to 1991 Salmon net pen Released 153
whale
Tasmania Bottlenose Tursiops truncatus 2 Prior to 1998 Atlantic salmon Fatal 60
dolphin sea cage
Tasmania Bottlenose Tursiops truncatus 3 1998–2000 Atlantic salmon Fatal 60
dolphin sea cage
Tasmania Short-beaked Delphinus delphis 6 1998–2003 Atlantic salmon Fatal 60,154
common sea cage
dolphin
Tasmania Dolphin Unidentified species 2 2012 Atlantic salmon Fatalg
sea cage
a
https://www.cowichanvalleycitizen.com/news/trapped-humpback-whale-freed-from-salmon-farm-near-tofino/.
b
https://salmonbusiness.com/whale-found-tangled-and-trapped-in-rope-dies-at-salmon-farm/.
c
https://www.stuff.co.nz/marlborough-express/news/69187299/dolphins-die-on-nz-king-salmon-farms.
d
https://www.fiskeridir.no/Akvakultur/Erfaringsbase/Knoelhval-i-merd.
e
https://www.cermaq.com/wps/wcm/connect/cermaq-no/cermaq-norway/baerekraft/asc-rapportering/.
f
https://www.fishfarmermagazine.com/news/whale-of-a-time-for-escaped-salmon/.
g
https://tassalgroup.com.au/wp-content/uploads/sites/2/2018/01/Tassal-Sustainability-Report-2012-13.pdf.

control nets set near Chile salmon farms are considered the principal 2015, a small minke whale broke into the cage and had to be eutha-
entanglement threat to cetaceans.110,164 In 2007, a Chilean dolphin nized to prevent further damages and fish escapes.69 In 2019, a 9-m-
140
was found entangled in salmon anti-sea lion nets and a humpback long minke whale broke through a cage, tearing a large hole in the net,
whale calf was fatally entangled in an antipredator net at a Chilean which enabled a small number of fish to swim free. The whale was
salmon farm141,142 (Table 4). An adult humpback whale was released released and the net was sealed but there was no further information
from entanglement in an antipredator net that was in the process of on the whale.152 The same news article mentions an interaction with
59
being installed at a salmon farm. Near the southern tip of Chile, a a minke whale at a different farm earlier that year but we were unable
15-m sei whale (Balaenoptera borealis Lesson, 1828) was believed to to confirm the report. Two humpback whales were entangled and
have died after it became entangled in a net at the salmon farm released from fish cages, one in 2009144 and a 9-m calf in 201569
(Table 3). The whale's body was almost entirely entangled with differ- (Table 3). The small calf swam into the net and its fluke became
165
ent length ropes and a metal chain was wrapped around its fluke. entangled in a rope, which made a small wound in the blubber. The
Chilean dolphins share coastal habitats with salmon farms and are farmers released the rope, attached another rope around the fluke,
most likely feeding on small schooling or benthic wild fish attracted to and pulled it out of the cage. The whale was released and observed
waste feed at the farms.59,66 In a study using predictive SDM, Hein- swimming with other whales shortly after the incident. A harbour por-
66
rich et al. partitioned fine-scale habitat use by Chilean dolphins and poise drowned after being entrapped at a salmon farm in 2018. At a
demonstrated potential overlap with mussel and salmon farms, with Scottish salmon farm in 2014, a juvenile male humpback whale
mussel farms showing higher probability of interactions. Because of drowned after being trapped under a net pen166,167 (Table 3). It was
the proximity of Chilean dolphin populations to farms,66 they may be noted that humpback whales are not usually observed around the
vulnerable to entanglements and entrapments; however, these events farms and the farmers surmised most likely the young whale was
are seldom recorded so it is not clear whether there are no interac- inquisitive and naïve.
tions or interactions are not documented. Marine finfish aquaculture has expanded in most Mediterranean
In Norway, minke and humpback whale interactions with Atlantic countries over the last two decades and depredation attempts by bot-
salmon farms have been documented. Minke whales have been tlenose dolphins attracted by farmed fish pose a risk of negative inter-
entangled in fish farm pens in Iceland and Norway. A minke whale actions. The antipredator net barriers used to protect the fish cages
was fatally entangled in an Iceland fish farm pen in 200567,143,144 from attacks by airborne and underwater predators present an entan-
(Table 3). In 2014, a 6-m minke whale swam through the upper part of glement risk. Observations of incidental catch of bottlenose dolphins
69
a net wall and into a cage. The fish farmers lowered a small part of in fish pens on the northeastern coast of Sardinia70 and of monk seals
the cage into the water, which created an opening for the whale to (Monachus monachus Hermann, 1779) in Turkey71,158 (Table 4) are the
swim out. No further damage to equipment or whale was observed. In result of inexperienced calves and loose predator barriers. A long-term
12 BATH ET AL.

TABLE 4 Documented accidental entanglements and entrapments of pinnipeds at marine finfish farms.

Species
Location common name Scientific name Number Year Gear type Outcome Citation(s)
South Australia Seal unspecified 2 2014–2016 Southern bluefin Released 155
tuna feedlot
South Australia Seals and sea lions unspecified 21 2014–2016 Yellowtail kingfish Released 155
sea cages
British California sea lion Zalophus 41 2011–2021 Net pen Fatal 139
Columbia californianus
Canada
British Harbour seal Phoca vitulina 50 2011–2021 Net pen Fatal 139
Columbia
Canada
British Northern fur seal Callorhinus ursinus 1 2021 Net pen Fatal 139
Columbia
Canada
Chile South American sea Otaria flavescens Net pen Fatal 59,60,156,157
lion
Chile South American fur Arctocephalus Net pen Fatal 60,157
seal australis
Faroe Islands Grey seal Halichoerus 10 2015–2019 Net pen Releaseda
gryphus
New Zealand New Zealand fur seal Arctocephalus 2 Prior to 2007 Net pen Fatal 43,56
forsteri
New Zealand New Zealand fur seal Arctocephalus 2 2014 Net pen Fatalb
forsteri
Tasmania Southern elephant Mirounga leonina 1 1998 Atlantic salmon Fatal 60
seal sea cage
Tasmania Leopard seal Hydrurga leptonyx 2 Prior to 1998 Atlantic salmon Fatal 60
sea cage
Tasmania New Zealand fur seal Arctocephalus 1 Prior to 1998 Atlantic salmon Fatal 60
forsteri sea cage
Tasmania Australian fur seal Arctocephalus 36 1998–2000 Atlantic salmon Fatal 60
pusillus sea cage
Tasmania Australian fur seal Arctocephalus 70 2010–2021 Atlantic salmon Fatalc
pusillus sea cage
Tasmania Seal Unspecified 18 2021 Atlantic salmon Fatald
cage
Turkey Monk seal Monachus 1 Fish farm Fatal 71,158
monachus
USA California California sea lion Zalophus 2 2005 Fish holding pen Fatal Jaclyn Taylor, NOAA
californianus attached to Fisheries, pers. comm.
dock 20 February 2022
USA Hawaii Hawaiian monk seal Monachus 1 2017 Marine fish cage Fatale Jaclyn Taylor, NOAA
schauinslandi Fisheries, pers. comm.
20 February 2022
USA Harbour seal Phoca vitulina 8 1996–1998 Net pen Fatal Jaclyn Taylor, NOAA
Washington Fisheries, pers. comm.
20 February 2022
USA California sea lion Zalophus 33 1996–1998 Net pen Fatal Jaclyn Taylor, NOAA
Washington californianus Fisheries, pers. comm.
20 February 2022
a
https://www.bakkafrost.com/media/1666/a71_y2016_w52.pdf.
b
https://www.stuff.co.nz/marlborough-express/news/69187299/dolphins-die-on-nz-king-salmon-farms.
c
http://tassalgroup.com.au/our-planet/reports/sustainability/.
d
https://dashboard.huonaqua.com.au/.
e
https://governor.hawaii.gov/newsroom/latest-news/dlnr-news-release-federal-and-state-agencies-investigate-death-of-hawaiian-monk-seal/.
BATH ET AL. 13

bottlenose dolphin distribution and behaviour study was conducted Off the coast of Hawaii farm workers at an almaco jack (Seriola
off the northeast coast of Sardinia, Italy, in the vicinity of a marine fish rivoliana Valenciennes, 1833) fish farm reported over 550 marine
farm with sea bass, gilthead sea bream, and meagre reared in 21 float- mammal observations from 2010 to 2016, and over 2500 from 2017
ing cages arranged in three rows of seven.73 In 2005, three animals to 2020.168,169 Bottlenose dolphins, humpback whales, and Hawaiian
were fatally entangled in 15 cm mesh antipredator nets (Table 3). Díaz monk seals (Monachus schauinsland Matschie, 1905) were frequently

pez and Shirai70 estimated one bottlenose dolphin fatal entangle-
Lo observed near the farm and in proximity to the cages. Individual ani-
ment per month in fish cages with loose antipredator netting and mals with distinguishing features were frequent visitors to the site but
zero for those with taut antipredator netting. During the entire study did not take up permanent residency. Pantropical spotted dolphins,
period (2004–2009), the estimated annual dolphin mortality was 1.5 rough-toothed dolphins (Steno bredanensis Cuvier, 1828), and false
per year based on five animals found entangled in nets.73 In the killer whales (Pseudorca crassidens Owen, 1846) have all been
Turkish Aegean Sea, farmers from 11 out of 25 surveyed sea bass observed in the area or in other offshore waters of the Kona Coast,
and sea bream fish farms reported individual monk seals were but have not been reported from the farm site.168 Dolphins observed
observed taking fish and damaging nets, mostly at night-time feed- at the farm site were reported to forage on wild fish, play, mate, fol-
ings during the winter months.71 A range of non-lethal deterrents low boats, and approach divers and cages (Jennica Hawkins, Ocean
was ineffective but antipredator nets were the only successful Era, pers. comm. 9 September 2021). Unfortunately, in 2017 at the
method to avoid fish loss. same farm, an endangered Hawaiian monk seal drowned in a partially
In Canada, public reports on authorised pinniped control activities decommissioned sea cage (Table 4). The 10-year-old male seal was
at British Columbia salmon farms are available on the Fisheries and discovered dead in a submerged empty fish cage in which farm
Oceans Canada (DFO) Public Reporting on Aquaculture - Pacific workers removed a side panel to allow a shark to swim out the previ-
Region database (DFO PAC-AQUA-MMI).139 From 2011 to 2016, ous day. Other than that incident, from 2010 to 2022, no entangle-
there were 249 authorised deaths of California sea lions (Zalophus ments, injuries, or mortalities occurred despite being located less than
californianus Lesson, 1828). From 2011 to 2015, 78 harbour seal one mile from the Hawaiian Islands Humpback Whale National
deaths were authorised, and two Steller sea lions (Eumetopias jubatus Marine Sanctuary. There was one fatality report of a juvenile North
Schreber, 1776) in 2011. The database also provides information Atlantic right whale (Eubalaena glacialis Müller, 1776) entangled in
about the numbers of known accidental marine mammal drownings at undefined aquaculture gear in the Western North Atlantic Ocean in
fish farms from 2011 to 2020. Animals often become tangled under- 2000.26,170 Although, the lack of standardised gear investigation pro-
water in the cage netting or other farm gear. During the time-period tocols at the time produced an incomplete investigation and there is
2011–2021, 41 California sea lion, 50 harbour seal, and 1 northern no direct evidence that aquaculture gear was involved (David Morin,
fur seal (Callorhinus ursinus Linnaeus, 1758) accidental drownings were NOAA Fisheries, pers. comm. 22 December 2020).
reported in Canadian Atlantic salmon fish farms (Table 4). Three dead Noteworthy, given anthropogenic and environmental threats to
humpback whales were discovered at Atlantic salmon farms and two sirenians worldwide,171–173 in the Philippines, local fishers have men-
trapped humpback whales were and successfully released. In 2013, a tioned instances of dugong (Dugong dugon Müller, 1776) entangle-
humpback whale was found dead at a farm but the necropsy report ment in the ropes of pearl farms and grouper culture cages.174 Aside
did not include the cause of death. In November 2016, two humpback from these anecdotes, we have not found validated documentation of
whales drowned in net pen gear; a juvenile breached the predator net manatee (Trichechus sp.) or dugong interactions with marine fish cage
of a net pen and the other was entangled by an anchor line, support- culture, yet at sites within their habitat range potential impacts to
ing an empty net pen.26,139 That same year, a third humpback whale these animals should be considered.55
was entangled at a fallowed farm and released alive. In 2018, a hump-
back whale was discovered swimming in an empty sea cage and not
entangled. Rescuers removed two panels from the antipredator net 3.1.3 | Underwater noise disturbances
and the whale swam out of the net pen139 (Table 3).
Review of the US NOAA Fisheries Marine Mammal Stock Assess- In addition to aquaculture farm construction and decommissioning,
ments database identified 41 unintentional pinniped fatalities: eight finfish and mussel farm operations produce underwater noise from
harbour seals and 33 California sea lions in Atlantic salmon net pens vessels, feeding systems, generators, aerators, net cleaning equip-
in Washington State from 1996 to February 2021 (Jaclyn Taylor, ment, and acoustic deterrents.175 There is evidence that underwater
NOAA Fisheries, pers. comm. 20 February 2022). In over 30 years of noise disturbances can alter the behaviour of marine mammals, cause
Atlantic salmon net pen farms operating in Puget Sound, Washington temporary or permanent injuries, or cause death, trigger a stress
State, there have not been any documented incidents of cetacean response, cause habitat displacement or avoidance, and disturb under-
entanglements in predator exclusion nets (Jaclyn Taylor, NOAA Fish- water acoustic cues for communication, navigation, and forag-
eries, pers. comm. 20 February 2022). In 2005, two California sea ing.43,68,175–182
lions were fatally entangled in predator nets of juvenile white seabass There are many factors that influence underwater noise, including
(Atractoscion nobilis Ayres, 1860) holding pens in the Channel Islands the number of pens, the operations on the farm, the characteristics of
off the California Harbour. the habitat, and the proximity to other sound-generating sources.
14 BATH ET AL.

Coastline configuration and habitat characteristics including water 3.1.5 | Attraction to artificial lighting
depth and sediment type impact noise sources and propagation.182
Whales may be more sensitive to increased noise production along Overhead lighting at fish farms provides navigational and personnel
migration routes,43,183–185 while pinnipeds demonstrate tolerance and safety, and farm security. Submerged artificial lighting is commonly
do not avoid underwater noises.179 Moreover, dolphins exhibit curios- used at higher latitudes to slow cultured fish maturation, increase
ity in response to underwater noises.43,186,187 Although not always growth rates, reduce fish densities near the surface, and evenly distrib-
effective on pinnipeds, the use of Acoustic Deterrent or Harassment ute the fish in cages.199–201 Overhead and submerged lighting around
Devices (ADDs and AHDs) to prevent pinniped predation has resulted finfish farms may attract marine mammals to caged prey or aggregated
in killer whale, harbour porpoise, and dolphin displacement from areas wild fish and cause trophic level disruptions.55,60,71,122,201–204 The
with active devices.43,175,176,178 Harcourt et al.188 tested the effec- effect of lighting on marine mammals is not only localised, as animals
tiveness of widely used commercial acoustic alarms to deter migrating may be attracted from longer distances, especially at night.163 In addi-
humpback whales from entanglement hazards including fish trap or tion, sound and light attract animals more than light alone. Light
pot lines. They detected no evidence of deterrence. The whales did shielded from all but essential directions minimises wild animal behav-
not speed up, slow down, or alter their course within the predicted iour disruption around finfish cages. Spotlights above pens positioned
audible range of the alarm. high above the surface will diffuse penetration through the water
column.201

3.1.4 | Vessel traffic

3.1.6 | Risk management and depredation
Depending on the size of the farm operation and the size of trans- mitigation
portation vessels, vessel traffic around aquaculture farms can pro-
duce noise, impose a navigation hazard, alter animal behaviour, Marine mammals can damage aquaculture gear, resulting in economic
exclude animals from habitats, and pose a collision risk. Vessel noise loss to the farm from equipment replacement costs and escaped fish
may contribute to stress and disrupt cetacean echolocation signals, from net breaches,62,71 and such interactions may cause serious injury
182,189,190
and thus reduce communication and foraging efficiency. or death to imperilled species. Predatory pinnipeds are often considered
Worldwide, vessel collisions with marine mammals have become a nuisance because of the net damage they incite by preying on the
recognised as a significant source of anthropogenic mortality and farmed fish.60,68,71,156,205–207 Torn nets not only allow for fish escapes,
191–195 194
serious injuries. Schoeman et al. found most scientific pub- resulting in economic loss for the farmers, but also present an additional
lications focus on collisions between large vessels and large whales. entanglement hazard to other animals. In addition, farmed fish are also
Their extended review discovered that at least 75 marine species are impacted by the stress-inducing predatory behaviour, which can indi-
vulnerable to vessel collisions, including smaller whales, dolphins, rectly effect fish growth and survival109 and result in lost biomass pro-
porpoises, dugongs, manatees, whale sharks, sharks, seals, sea otters, duction. Thus, deterring pinnipeds from salmon net pens in the United
sea turtles, penguins, and fish. States,62,208 Canada,63 Scotland,103,209 Australia,210 and Chile141,156,163
While marine mammal behaviour changes in response to vessel has been a constant battle for decades.109 Until recently, authorised kills
181,196,197
traffic have been studied, documented cases of direct vessel have not only been legal but also tolerated as an effective control mea-
traffic effects on protected species around aquaculture farms are diffi- sure by salmon farmers in some countries62,63,68,109; although, reliable,
cult to find. Vessel traffic related to aquaculture affects Chilean dol- quantitative mortality data is lacking.109 An extensive review of pinniped
phins by altering behavioural responses such as changes in swimming interactions with salmon farms in Canada, published by Jamieson and
198
reorientation rate and speed. While individual farms may have very Olesiuk,63 described non-lethal intervention methods, the effects of
little vessel traffic, areas with multiple farms may have a cumulative lethal deterrents to sea lion and seal populations, as well as the financial
effect. The impacts of vessel noise and collision impacts on blue impacts to the industry. Although farm stock or gear damage may be a
whales (Balaenoptera musculus Linnaeus, 1758) in northern Chilean few thousand dollars for an individual farm, multiple cases can amount
Patagonia is significant considering the aquaculture fleet is an order of to millions of dollars in 1 year for a country. The growth of the fish farm-
magnitude larger than any other sector including cargo, transport, arti- ing industry and concurrent pinniped population expansion has
sanal fishery, and industrial fishery fleets.195 Using density predictions increased the number of interactions, and previously accepted lethal
from previous SDM, spatially explicit predictions of behavioural control methods are less viable due to ecosystem conservation objec-
responses to vessel presence, and vessel tracking data, they estimated tives and regulatory protection. In many cases, public support for aqua-
the relative probability of vessels encountering whales and identify culture is decreased when there is news of marine animals being
areas where interaction is likely to occur. Given the size of finfish and harmed or culled.
mussel culture operations in Chile, the projected industry growth,115 To reduce marine mammal depredation or avoid injuries and
and known presence of several marine mammal species, vulnerable lethal interactions, aquaculture operations use a suite of methods:
protected species are at risk of collision with vessels associated with harassment, aversive conditioning, non-lethal removal, lethal removal,
aquaculture activities. population control, and exclusion.5,41,71,109,209,211,212 Harassment by
BATH ET AL. 15

chasing, explosives, and ADDs is effective in the short term but tend Acoustic Startle Technology reduces the avoidance response to a
to be less efficacious over time as animals become acclimated to the defined area and decreases noise pollution.223
60
noise. ADDs emit sound underwater at a range of frequencies to In addition to the ADDs, Terhune et al.211 reviewed other non-
deter predation on the stocked fish by causing auditory discomfort. lethal interventions or aversive conditioning such as harassment by
Noise harassment devices may actually become attractants to habitu- boat or with noise (such as underwater seal firecrackers), predator
ated individuals who associate the unpleasant sound with an easy models or sounds, acoustic devices, and relocation. Seal bombs and
43,186,187
prey source over time. The effectiveness of noise deterrents shooting are most effective if used before animals acclimated to fin-
and their secondary impacts to non-target animals is uncer- fish cages and established a permanent interest in the farm.224,225
68,163,178,211,213,214
tain. In a controlled experiment to assess the Physical predator models and sound devices (imitating killer whales,
influence of an ADD on free-range bottlenose dolphin behaviour by for example) are also not effective.60,103,156 Electric fences have been

pez and Mariño,
Díaz-Lo 215
dolphins were not deterred from the farm, used to prevent pinnipeds from hauling out on farm structures in the
especially when food was present. ADDs and underwater explosive Canada and the US Pacific Northwest.48,224,226 Capture and reloca-
devices used to prevent pinniped predation have resulted in killer tion of destructive animals is time-consuming, expensive, and mini-
whale, minke whale, harbour porpoise, and dolphin displacement from mally effective.211
43,103,176,178,216–220
areas with active devices. Until recently, pinniped lethal removal from aquaculture farms
In Scotland and British Columbia, Canada, harbour porpoises was common practice and authorised in many countries with large-
avoided salmon farms when ADDs were active but returned quickly scale Atlantic salmon farms, including Canada, Chile, Norway, and
when they were deactivated.176,178 In farms where active ADDs had Scotland. Before the spring of 2020, Fisheries and Oceans Canada
been deployed for some time, animals were observed foraging. In (DFO) authorised farm licence holders227 to lethally remove nuisance
New Zealand, Hector's dolphins, avoided acoustic gillnet pingers, and seals that posed eminent danger to the aquaculture facility or to
based on this observation, Stone et al.221 suggested using similar human life, if all other non-lethal deterrent efforts failed. In Chile, all
devices at salmon farms to deter pinnipeds could also impact non- marine mammals are protected by law from intentional killing and
target mammals. Killer whales in British Columbia avoided marine accidental mortalities in fishing operations are legally required to be
farm areas where ADDs are in use216,222 but whales in a nearby farm reported.228 Despite this law, strict adherence to the regulation and
without ADDs remained stable during this same period. Six years after accountability is lacking. In 2021, rules governing the interaction
deployment, the local killer whale abundance returned to previous between marine mammals and aquaculture require antipredator nets
216
levels after the devices were removed. In the Bay of Fundy, ADDs be installed at salmon farms and detailed plans for dealing with sea
deployed near aquaculture facilities did not elicit startle responses, lions must be included in farm operational management plans.229 In
cause measurable avoidance behaviour, or change haul-out behaviour Scotland, shooting seals was licenced to aquaculture operations until
of pinnipeds over many years.211 Salmon farm managers surveyed in June 2020 when the Scottish Parliament approved an amendment to
Scotland questioned the effectiveness of ADDs and do not use them the Marine Scotland Act of 2010,230 banning intentional killing of
178,209
at all farms. Despite daily sightings of seals near farms, the seals to protect cultured animals.231 Animal welfare concerns have
authors suggested seal predation on farmed fish decreased over the driven these changes. Importantly, under the US seafood import
previous decade and less than a quarter of salmon farms reported rule,232 exporting countries must implement a regulatory program com-
major problems with seals. Because specific individuals allegedly were parable in effectiveness to US policy. Beginning in January 2023, such
responsible for the most damage, improving individual recognition programs must be implemented in which intentional marine mammal
techniques was a priority for farm management and reduced interac- kills are prohibited, marine mammal population assessments include
tions. In a review of commercial ADDs, Götz and Janik213 concluded bycatch estimates, bycatch limits are quantified, and bycatch reduction
main problems were reduced efficiency over time and noise pollution measures are implemented.59,232,233 Because the United States is a
that impacted communication, hearing—including permanent damage, major importer of Atlantic salmon from Canada, Chile, Norway, and
and behaviour of non-targeted animals where ADDs were employed. Scotland, documenting marine mammal interactions at farms is now
They also suggested ADDs used to deter pinnipeds from farms oper- imperative to comply with US seafood import requirements.59,234
ate at the same frequencies received by odontocetes with more acute Under this import rule, exporting countries must implement a regula-
hearing sensitivity and may explain large-scale habitat exclusion in the tory program in which intentional marine mammal kills are prohibited,
vicinity of these units. Findlay et al.217 quantified the cumulative stock assessments are conducted, bycatch estimates are quantified,
impact of multiple farms using ADDs along the Scottish west coast and bycatch limits are imposed.232
and demonstrated ADDs are a significant and chronic source of Other farm management practices including the addition of false
underwater noise disturbance. Using species-specific frequency- bottoms to avoid predation under the cages, increased net tension,
dependent hearing sensitivity, deterrent devices can be modified to removal of fish carcasses, and installing antipredator nets are accepted
target one taxon and minimise impact to others. Götz and Janik223 tar- as best practices. Advances in antipredator cage technology, routine
geted grey seal acoustic startle reflex with transducers that emitted net maintenance to avoid fish escapes, minimal use of lights at night,
short, isolated noise pulses at low duty cycles, and elicited avoidance and feed management to reduce waste are farm practices that likely
responses by the animals within 250 m of the units. This Targeted decrease the potential for negative interactions. Successful pinniped
16 BATH ET AL.

deterrence is achieved using physical exclusionary barriers, including turtles and cheloniid turtles behave differently in response to lines used
rigid net materials for fish cages or the installation of rigid exclusion- in commercial fishery gear,237,238 and thus, the vulnerability and
ary nets around finfish farms to reduce injury and siting cages off- mechanics of entanglement are likely different among mussel farm gear
shore, away from haul-out sites and rookeries.48,55,62,71,114 Exclusion types. Uncertainty remains about the entanglement risk posed by high-
nets must be strong enough to resist chewing or tearing and should tension horizontal backbone lines at mussel longline farms because
be properly tensioned to prevent entanglement.25,114 Innovative net there have been no published reports of entanglements. While ten-
materials, like the Fortress Pen, patented by Huon Aquaculture and sioned lines may decrease entanglement risk for small cetaceans and
made from Kevlar and woven nylon, the same material as bulletproof cheloniid sea turtles, there is still concern that large whales and leather-
vests, provides high net visibility and a robust barrier to seal entry. back turtles may be at risk for entanglement and/or injury if they collide
Farms located distant (>20 km) from haul-out sites tended to have with these lines at mussel farms.
55
fewer pinnipeds trying to forage on farmed fish. In South Australia, There are three known incidents involving leatherback sea turtles
predation by New Zealand fur seals and Australian sea lions (Neophoca entangled in mussel ropes in Notre Dame Bay, Newfoundland, Canada
cinerea Péron, 1816) at southern bluefin tuna farms was considered a (Table 5). In 2009, one turtle was found dead and rolled up in the mus-
continuing problem for tuna farmers in Port Lincoln, causing a signifi- sel farm lines.237,245 Two individuals were reported entangled in mussel
cant cost to the industry.153,235 Fur seals were most commonly seen spat collection lines, one leatherback was found dead at depth in 2010.
around the farms and in the cages, but the sea lions were aggressive In 2013 the second was found alive at the surface and released after
predators that also stressed and damaged the tuna. The fur seals were being disentangled around the head and flippers.67,237,246 One leather-
too small to be a threat to the tuna and fed mostly on the tuna feed back was entangled in vertical line anchoring gear associated with a
and other small fish around the cages. Although fencing was the best shellfish aquaculture site in the Greater Atlantic Region in US waters in
method used to deter seals, the most frequent entry method to the 2014 and released26 (Kate Sampson, NOAA Fisheries, pers. comm.
tuna cages for seals was jumping over the fences. Frequent cage NOAA NMFS Sea Turtle Disentanglement Network database [STDN];
maintenance, hole repairs, and tuna carcass removal was effective in Table 5). On June 11, 2022, a 900-lb leatherback was entangled off
235
minimising seal and sea lion attacks. Nantucket, Massachusetts, USA in research aquaculture gear for bay
scallop spat collection (David Morin, NOAA Fisheries, pers. comm.
20 July 2022; Table 5). The turtle was found in 3 feet of water
3.2 | Sea turtles entangled around the neck twice and both flippers with two complete
sets of gear including four cinder blocks, six surface buoys, and 48 m of
There are seven species of sea turtles worldwide, including green (Chelo- line. The turtle was completely disentangled and released alive. At an
nia mydas Brongniart, 1800), hawksbill (Eretmochelys imbricata Fitzinger, almaco jack farm in Hawaii, sea turtles have not been observed around
1843), Kemp's ridley (Lepidochelys kempii Garman, 1880), leatherback the fish cages. Because green sea turtles are common in the nearshore
(Dermochelys coriacea Vandelli, 1761), olive ridley (Lepidochelys olivacea waters of the main Hawaiian Islands, it is likely they occasionally swim
Eschscholtz, 1829), loggerhead (Caretta caretta Linnaeus, 1758), and through the farm area despite no recorded sightings.168 Additional
flatback (Natator depressus Garman, 1880). All species except for flat- records of sea turtle and aquaculture farm interactions were not con-
back, which is data deficient, are classified as vulnerable, endangered, or firmed in this investigation. In Chile and the Philippines, turtles interact
236
critically endangered. To date, there are few reported incidents of with aquaculture farms, primarily mussel, pearl, and seaweed, but are
sea turtle injuries or mortalities at aquaculture sites.237,238 Interactions considered a nuisance and therefore are harvested, intentionally killed,
have been reported at mussel farms in Newfoundland, Canada, and and are not protected or reported.59,137 Aquaculture gear impacts to
Chile, and pearl and seaweed farms in the Philippines. Sea turtles are sea turtles are understudied and need to be considered in future entan-
observed as incidental visitors around marine fish cages but not per- glement and mitigation actions.237 Entanglement injuries can result in
239,240
ceived as predatory threats to the farmed fish. Because they are reduced feeding efficiency, impaired locomotion, exertional myopathy,
protected in many countries as threatened or endangered species, the compromised blood flow and necrosis, infections, and prolonged, debili-
primary concern with sea turtles is the threat of entanglement with nets, tating health complications or death.237,247
lines, or other floating equipment at aquaculture farms, although vessel In the United States, NOAA Fisheries held a workshop in 2008 to
traffic around farms could also cause collisions.238,241–244 Relatively little address sea turtle species distribution in the Northeast US, interac-
is known about how sea turtles may be impacted by marine fish cage tions with vertical lines in fixed gear commercial fisheries, injury
farms and after an exhaustive search, we were unable to find published assessments, and disentanglement techniques.248 The participants
reports of harmful interactions. suggested some sea turtles become entangled in vertical lines by
chance, because of general curiosity, or as they forage. Participants
agreed that fishing gear is often set in areas where turtles forage and
3.2.1 | Entanglement thus present an opportunity for entanglement. In an analysis of leath-
erback turtle interactions with fixed gear fisheries over almost two
Sea turtle entanglement reports at aquaculture farms are rare; however, decades of observations, Hamelin et al.237 determined turtles are vul-
from commercial fishery gear observations, they are vulnerable to nerable to both horizontal and vertical lines. Slack lines pose the
entanglement in both horizontal and vertical lines.237 Leatherback greatest entanglement threat because the lines wrap tightly, multiple
BATH ET AL. 17

TABLE 5 Documented sea turtle entanglements at shellfish farms.

Species
Location common name Scientific name Number Year Gear type Outcome Citation(s)
Newfoundland Leatherback sea Dermochelys 1 2009 Mussel farm Fatal 237,245
Canada turtle coriacea
Newfoundland Leatherback sea Dermochelys 1 2010 Mussel farm spat Fatal 67,237
Canada turtle coriacea line
Newfoundland Leatherback sea Dermochelys 1 2013 Mussel farm spat Released 67,237,246
Canada turtle coriacea line
USA Greater Leatherback sea Dermochelys 1 2014 Shellfish farm Released NOAA Fisheries STDN,
Atlantic Region turtle coriacea vertical anchor unpublished dataa
line
USA Nantucket, Leatherback sea Dermochelys 1 2022 Bay scallop spat Released D. Morin, NOAA Fisheries,
Massachusetts turtle coriacea collection gear pers. comm. 20 July
2022
a
NOAA Fisheries Sea Turtle Disentanglement Network (STDN) database.

times around flippers as turtles try to escape. Because they are unable turtles are most susceptible to vessel strikes when they surface to
to free themselves, they can drown if held under water and their sur- breathe, feed, bask, mate, and orient themselves to their surround-
vival depends on human intervention to remove the lines.237 These ings.255 Sea turtles rely primarily on visual cues to detect vessels256
conclusions are validated in an analysis of 15 years of entanglement and may have limited ability to manoeuvre in the water column to
reports in fixed gear fisheries off Massachusetts, USA.238 The leather- avoid collisions, depending on the vessel size and speed. Inclement
back entanglements at mussel farms occurred in spat collector lines weather and reduced light at night decreases visual acuity and
that are typically not anchored to the substrate. Fixed gear that increases the collision risk. All seven species of sea turtles have been
is highly weighted may pose an immediate threat to leatherbacks injured or killed by vessel strikes.194 It is unknown what proportion of
when they are entangled at depth because they cannot surface to sea turtles struck by vessels survive their injuries. It is difficult to
237,238
breathe. quantify the impact of vessel strikes on sea turtles because their bod-
ies are negatively buoyant and sink, so fatal vessel collisions may go
undetected and unreported. There is good evidence from sea turtle
3.2.2 | Underwater noise disturbance stranding data from Florida, USA, that postmortem vessel strikes are
rare and the vast majority of stranded animals with vessel strike inju-
Underwater noise disturbances from aquaculture farm construction, ries were hit antemortem and died as a result.257 Schoeman et al.194
operations, or decommissioning could potentially alter the environmen- provide detailed mitigation measures to improve vessel safety around
tal soundscape and impact sea turtles in the vicinity of these activities. marine animals to prevent collisions.
While underwater noise impacts to marine mammals are well-
studied,177,179–182 impacts to sea turtles are relatively uncertain.249,250
It had long been assumed sea turtles do not vocalise and little is known 3.2.4 | Risk management
how sea turtles use auditory cues to avoid predators, locate prey, or
navigate their environment.251,252 Experimental studies have verified Given the potential significant impact of entanglement on sea turtles in
sea turtles can detect sounds both in air and underwater253 and recent aquaculture gear, detailed accounts at mussel and finfish farms are par-
research on green sea turtles by Charrier et al.252 suggests acoustic amount to understand the frequency and severity of encounters. To
intra-specific communication may exist. Sea turtles inhabit different reduce negative interactions, best farm management practices include
254
ocean habitats throughout their life history. Juveniles and adults the use of rigid netting material for the cage, keeping mooring lines
spend most of their time in the inshore environment, which is typically taut, and removing any loose lines or floating equipment around the
noisier than the open ocean pelagic habitat where hatchlings feed and farm. Building on the 2008 NOAA Fisheries sea turtle workshop,248
grow. The location of the underwater noises from farms and associated NOAA Fisheries and a representative from Fisheries and Oceans
vessels could differentially impact important sea turtle behavioural and Canada met to develop a summary of relevant information and a matrix
ecological functions. of gear research ideas.258 Many fisheries gear modifications presented
in the matrix to minimise turtle entanglements provide possible modifi-
cations to vertical lines used in aquaculture and could be explored fur-
3.2.3 | Vessel traffic ther. In addition to gear modifications, minimising marine debris
reduces the potential for turtles to ingest trash associated with farm
In addition to underwater noise, vessel traffic around aquaculture operations. Because ADDs used to deter marine mammals are outside
farms can create a navigation hazard and pose a collision risk. Sea the frequency turtles detect underwater,251,259 they would likely not
18 BATH ET AL.

be effective at keeping sea turtles away from gear. However, low fre- of the physical structures of mussel and finfish farms,36,56,262 and the
quency acoustic alerts targeting sea turtle sensitivity ranges, have been presence of added activity and boat traffic around these farms dis-
used in gillnet fisheries in Baja, Mexico and showed a 60% reduction in rupts seabird breeding and feeding behaviour.262,263
turtle bycatch per unit effort (Dow Piniak, NOAA Fisheries, pers. comm. A study in southwest Ireland89 found no adverse effects on seabird
238
9 May 2022). Dodge et al. recommended the need for maintaining a species richness or overall abundance at nearshore mussel farms
trained and active disentanglement network for bycatch mitigation, (14–17 m deep). More birds, namely cormorants and gulls, were present
which is critical not only for turtles, but marine mammals as well. Coor- in mussel farm areas. The farm provided structure for perching and
dination with these disentanglement networks is paramount for marine source of food from epifauna growing on above-water structures. Nei-
aquaculture farms where vulnerable species are present. ther benefit would be expected at submerged offshore mussel farms,
which do not include rigid structures suitable for perching. In a compari-
son of seabird activity budgets between three areas of nearshore mussel
3.3 | Seabirds farms and three control sites in Bantry Bay, Ireland, Roycroft et al.265
concluded the impact of mussel suspension culture appeared to be posi-
Marine aquaculture farms across different regions may interact with tive or neutral on seabirds at the study site. Aguado-Giménez et al.264
species of albatross, cormorants, gannet, loons, pelicans, auks, gulls, observed the spatiotemporal variability of piscivorous sea birds over a
petrels, storm petrels, shearwaters, diving ducks, penguins, and terns, year at eight fish farms in the western Mediterranean and recorded sea-
among others. Depending on the geographic range of these seabird spe- sonal differences in bird density and assemblages. Bird density increased
cies, the potential for overlap with aquaculture farms is an imperative from fall to winter and decreased in spring and summer, which was par-
260
siting consideration. Both mussel farms and finfish farms attract tially explained by season and distance to breeding/wintering grounds. In
seabirds.36,109,261–264 Marine aquaculture farms may exclude seabirds Admiralty Bay, New Zealand, Fisher and Boren273 surveyed king shag
from important habitats including migratory routes and feeding grounds, (Leucocarbo carunculatus Gmelin, 1789) foraging distribution and habitat
or may cause benthic disturbances that cause high turbidity and reduce use around mussel farms. Although a few observations of this behaviour
foraging success, altered prey availability, foreign object ingestion, have been reported, they did not observe birds foraging in the farms.
entanglement, and collision with farm structures.261,263,265,266 Distur- The birds perched on farm structures to rest, roost, and preen. In Chile,
bances to breeding colonies may result in nest abandonment, reduced Jiménez et al.274 observed higher seabird abundance at salmon pens
breeding success, and localised population depletion.263,267 compared with their control sites and found salmon farming had no sig-
nificant effect on avian species richness. The primary species included
diving piscivores, perching piscivores, omnivores, and carrion eaters but
3.3.1 | Habitat modification, attraction, or exclusion did not include herbivores, invertebrate, or surface feeders.

The attraction and aggregation of forage fish around farm structures

as well as the farmed mussels and net fouling organisms can 3.3.2 | Entanglement
provide enhanced feeding opportunities for marine diving
birds.58,114,224,225,261,263,266 In Washington State, diving birds feed on Entanglement poses the biggest threat to seabirds in both mussel42,275
239,268
colonising epifauna that accumulates on farm structures. Envi- and finfish aquaculture operations48,68,262; however, entanglement data
ronmental changes to benthic communities in proximity to fish farms resulting in injury or mortality are rarely available. Seabirds are at risk of
could potentially disrupt diving bird feeding preferences.266 Flocks of becoming entangled in lines or nets, colliding with structures while flying,
diving ducks such as the black scoter (Melanitta nigra Linnaeus, 1758) and ingesting debris, all of which may result in injuries or death.25,56,261,262
or common eider (Somateria mollissima Linnaeus, 1758) can substan- Ingestion and entanglement of marine debris from associated farm activi-
tially impact mussel biomass by predation at farms.30,224 Small fish ties could block seabird digestive tracts and cause serious injury or
commonly consumed by birds are attracted to net pen structures, death.261,262,276,277 In 2003, Lloyd36 reported there were no published
residual feed, and farm lighting.204,269,270 Seabirds are also attracted accounts of seabird entanglements in New Zealand aquaculture. More
to lights around net pen facilities, which put them at risk of colli- recently, both Huon Aquaculture162 and Tassal Group278 report bird
261–263,271
sion. Increased bird populations around farm structures releases and mortalities at their Tasmanian salmon farms in real time on
may add large amounts of nutrients to the surrounding water, affect- their sustainability dashboards.
ing fouling organisms on nets.114,261 Adding nutrients to the water Incidental seabird bycatch during commercial fishing operations is
column can also cause algal blooms and alter birds' ability to locate recognised as a global problem, but there are no official reports of
prey due to increased water turbidity. seabird deaths as a result of entanglement in fixed lines of the type
Floating farm structures may provide roosting locations close to found in mussel farms or spat catching areas.276,277 In Marlborough
foraging grounds and afford protection from terrestrial preda- Sound, New Zealand, adult and young Australian gannet (Morus serra-
56,262,266,272
tors. However, the refuge and increase in foraging effi- tor GR Gray, 1843) have been found entangled in rope ties from mus-
ciency could also alter foraging behaviour, resulting in changes in the sel farms incorporated into their nests.36,272 At finfish farms, diving
food web or disturb breeding colonies. Habitats available for surface birds become entangled in underwater fish containment nets and
feeding birds (gulls, terns, shearwaters) can also be reduced because drown. Predator exclusion nets can also entangle birds, resulting in
BATH ET AL. 19

limb injuries or death. Seabirds are considered a low predatory risk to Management of Sharks283 provides guidance for member nations to
the live farmed fish but may scavenge mortalities or take fish during develop their own shark conservation management plans. Under the
21,239,242,268
transfer or harvest. Convention on Migratory Species intergovernmental treaty, 49 Mem-
ber States and 12 Cooperating Partners signed a Memorandum of
Understanding on the Conservation of Migratory Sharks in 2010,
3.3.3 | Risk management amended in 2018 (https://www.cms.int/sharks/en).284 Backed by the
United Nations Environmental Program, this memorandum is a legally
Richman279 recommended deterrent methods to reduce sea duck non-binding international instrument. The agreement is to achieve
depredation at mussel farms. She noted, although loud sounds and maintain a conservation status for migratory sharks based on the
frighten birds, they can become desensitised and habituated. Visual best available scientific information, considering the listed species'
devices like streamers, reflective mirrors, and model predators, are socioeconomic value in various countries. Given the recent global
minimally effective. Human activities, boat chasing, and falconry are interest in shark population declines and the need to implement con-
effective but labour intensive. Exclusionary nets are effective for small servation efforts,285–288 the potential impacts of offshore aquaculture
nearshore mussel farm sites but less practical for large offshore farms. to sharks are important to evaluate.26,147,289,290
And while shooting is highly effective at the individual level, it Aggregating fish around farms attract sharks to finfish farms in
requires permits and may be socially unacceptable. Puerto Rico,291 Hawaii,292 the Bahamas,293 Canary Islands,294 Latin
Net material and size play an important role in entanglement risk. America,226 the US Pacific Northwest,239 New Zealand,56,290 and
Nets with large meshes, small diameter twine, or transparent monofil- Australia.25,147,289,295 In addition to wild fish attracted to farms, sharks
280
ament are more likely to cause seabird entanglement. Fortunately, are likely attracted to multiple stimuli associated with fish farms,
offshore marine fish farms do not require small diameter, transparent including live fish in cages, the presence of dead fish at the bottom of
nets for any component of the cage design and farming operation. cages, the odour trail generated during feeding, farming operation
Research conducted by Nemtzov and Olsvig-Whittaker281 examined sounds, and the physical structures.58,147,289,290 Sharks damage farm
101 netted marine cages using 11 different net types varying in mesh structures and cause economic impacts to the farms through fish
size, material, colour, and thickness. They studied the influence these escapes and predation as well as decreased production from cultured
net design features had on bird mortality. Bird mortality was largely a fish under regular attack.151
function of net visibility.281 Fewer birds were entangled in nets with
dark-coloured netting, meshes ranging from 20 to 30 mm, and made
of woven nylon 1.8–2.0-mm thick. 3.4.1 | Shark presence at offshore farms
Enclosing predator nets at the bottom of cages and using top nets
over cages to exclude birds, ensuring nets are kept taut, and frequent Four species of sharks frequent New Zealand King Salmon farms
maintenance decrease the number of entanglements and subsequent including spiny dogfish (Squalus acanthias Linnaeus, 1758), bronze
mortality in exclusion nets.261,280,281 Exclusion nets can be an effective whalers (Carcharinus brachyrus Günther, 1870), blue sharks (Prionace
solution; however, they cannot be used everywhere and thus must be glauca Linnaeus, 1758), and seven-gill sharks (Notorynchus cepedianus
used discriminatingly.280 Any change in cage design must consider the Péron, 1807) because they are primarily attracted to dead fish at the
153
method by which avian predators forage. Net pen site selection to bottom of the cages.290 In Australia tuna lots and yellowtail kingfish
avoid overlap with home ranges, critical breeding grounds, and foraging and mulloway (Argyrosomus japonicas Temminck & Schlegel, 1843)
habitats is prudent.241,244,261,262 Siting farms in areas with strong currents cages, interactions with pack-hunting bronze whalers are a more sig-
to disperse nutrients away from fish cages and minimising feed waste nificant issue than opportunistic, solitary hunting white sharks (Carch-
reduce the potential for negative interactions fostered by aggregating arodon carcharias Linnaeus, 1758); however, white sharks attract more
prey.261,262 The light type and direction are important to abate collisions attention because they are a higher profile species and protected
with net pen structures that could result in injury or death.261,262 under the Environment Protection and Biodiversity Conservation Act
Curtailing the growth of biofouling on nets and keeping them taut of 1999.289 According to the Tuna Boat Owners' Association in
reduces the risk of attraction and entanglement. Debris removal and con- Australia, interactions with white sharks and bronze whalers tend to
tinuous monitoring diminish the risk of ingestion or entanglement.261,262 occur in specific areas and at individual farms, during fish transfer
It may be possible to reduce the negative impacts caused by human dis- from towing to farm pontoons in the grow-out cycle, and in seasonal
turbances at seabird breeding grounds by situating aquaculture farms patterns.289
263
away from seabird colonies and nearby foraging areas. In the Mediterranean Sea, white sharks have been sighted near
tuna farms. In Norway, spiny dogfish are attracted to dead fish in the
bottom of salmon cages.290 A telemetry study of sandbar (Carcharhi-
3.4 | Sharks nus plumbeus Nardo, 1827) and tiger sharks (Galeocerdo cuvier Péron &
Lesueur, 1822) near almaco jack cages off Hawaii found sharks aggre-
Consistent with the FAO Code of Conduct for Responsible gated near the cages with some individuals recorded for the 2.5-year
Fisheries,282 the International Plan of Action for the Conservation and study.168,292 From June to August 2008, shark bites of various sizes
20 BATH ET AL.

were discovered in the webbing of one cage and were immediately minimal threat to the farm workers. Another time, a humpback
repaired. In 2009, a small Galapagos shark (Carcharhinus galapagensis whale carcass floated near the farm, drawing a swarm of tiger
Snodgrass & Heller, 1905) breached the cage and was caught and sharks around the fish cage. The farm workers were able to drag
released unharmed. In response to these net breaches, the farm the dead whale offshore so it would not wash up on the beaches or
changed the cage mesh material to deter sharks and prevent further pose risk to the farm operations (Jennica Hawkins, Ocean Era, pers.
damages.168 Off Réunion Island, video monitoring documented 190 bull comm. 1 December 2020).
shark (Carcharhinus leucas Müller & Henle, 1839) observations over
1 month under a sea cage fish farm. Many sharks were re-sighted and
at least three individuals displayed site attachment throughout the 3.4.3 | Dietary shift
month-long study.296 The authors reported that no sharks were
observed attacking the cage to access the farmed fish. Sharks are strict carnivores and consume a wide range of prey
including plankton, invertebrates, teleosts, elasmobranchs, birds, rep-
tiles, and marine mammals. In most shark species, prey type and size
3.4.2 | Threats to gear and workers may change with shark ontogenetic shifts, expanded ranges, and
improved hunting skills.301,302 Generally, sharks feed opportunisti-
In Norway, spiny dogfish bite through holes to prey on caged salmon cally on the most abundant prey item, primarily fish.302,303 Sharks
151
and are a well-known challenge for fish farmers in some areas. In can be attracted to fish farms because wild fish often aggregate
southeastern Tasmania, large sharks were recorded twice as the cause around the farm structures.8,56,61,122,269,270 The nutritional value of
of large (1.5 m) tears in fish farm containment nets; one of these these wild fish could change if their diets primarily consist of unea-
attacks was by a thresher shark (Alopias vulpinus Bonnaterre, ten feed pellets from farmed fish, as compared with their natural
1788).153 Yellowtail kingfish farms in Australia are frequented by prey sources. Although many aquafeeds are most commonly com-
bronze whalers, which tend to aggregate in groups of 4–14 fish. The posed of fishmeal and fish oil, many manufacturers are moving to
sharks typically break through the bottom of the nets but have also more sustainable plant-based ingredients, which modify the fatty
broken through the sidewalls.289 In South Australia, aquaculture cages acid composition and fat content levels of tissues of wild fish that
have been identified as an entanglement threat to white sharks.297 feed on the lost pellets.6,304 In addition, antibiotics in some feeds
The sharks will break into aquaculture cages in search of food, posing have the potential to accumulate in the tissues of the wild fish and
295
a risk to the farmed stock and cage operators. result in liver damage, acute toxic effects, bacterial resistance, and
Increased encounters with sharks could occur if the animals are immune system suppression.298
attracted to net pens and share space with recreational or commer- Octopus, molluscs, and crustaceans are prey for small bottom-
cial divers.298 Because of these threats, dangerous species are dwelling and bottom-feeding sharks.302 Benthic macro-invertebrate
sometimes lethally removed from marine farm areas. For safety rea- communities can change in the presence of aquaculture farms
sons, some farm managers have killed sharks before removing them because of crop and biofouling organism drop-off and the deposition
from cages. Before 2001, within a 5-year period, there were nine of farmed fish faeces and uneaten food pellets.56,305–308 If preferred
confirmed white sharks captured in tuna ranches, of which six were prey items are not available, sharks may be forced to alter their diets
299
killed and three were already dead. In Australia, an estimated and scavenge dead fish accumulating in cages and uneaten feed
20 white sharks a year are killed at marine aquaculture farms,295,299 beneath farms.289,290,298 Depending on the shark species and their
and in response to the need for better conservation practices, live- foraging habits, these farm-induced dietary changes raise concern
release methods have been developed in South Australia.289 In about the potential effect of farms on the biology of these predators,
New Zealand, culling in and around farms happens infrequently and their trophic interactions, and ecosystem function at different spatial
according to anecdotal information, shark mortalities from entan- and temporal scales.
glement or entrapment are rare.290 In South Africa, a salmon farm
situated inside an ecologically significant white shark congregation
area closed after eliciting major negative public reactions.300 In the 3.4.4 | Risk management
first year of operations (2005) at a fish farm in Hawaii, cage divers
were not removing all moribund fish from the cages. An aggressive As the marine aquaculture industry moves into offshore sites, the
tiger shark took up residency at the farm to feed on the dead fish potential economic and ecological risk of large-scale fish releases due
and displayed agonistic behaviour causing divers to exit the water to sharks tearing nets may be a concern depending on net types and
for safety. To protect the employees from future attacks, the shark locations used.8 Technological improvements in aquaculture cage
was killed. Because using this long-term mitigation strategy was materials like strong PET monofilament and semi-rigid and tightly
unacceptable, the farm worked in collaboration with state agencies stretched net walls make cultured fish resistant to predator attacks.47
to develop a shark management plan, which included relocation, as The semi-rigid net structure is designed with self-closing properties
recommended by catch and release research.168 After that incident, that prevent fish escape and thermo-formed double twisted mesh to
the tiger and sandbar sharks observed around the farm posed prevent unravelling. The degree to which sharks are attracted to farms
BATH ET AL. 21

to feed on wild fish, farmed fish, or sunken pellets, and the resulting 4 | C O N C LU D I N G R E M A R K S
behavioural or ecological impacts are unknown. Although farming off-
shore may reduce exposure to coastal predators such as pinnipeds The growth of coastal and offshore aquaculture worldwide is drawing
and coastal cetaceans, exposure to predatory sharks may increase. In attention to the potential environmental impacts, including impacts to
addition, sharks that spend a significant amount of time close to the protected species. Despite an increased understanding of how pro-
surface like whale sharks (Rhincodon typus Smith, 1828) and basking tected species may be affected by marine aquaculture farms over the
sharks (Cetorhinus maximus Blainville, 1816) may be more vulnerable last decade, questions remain. The research and data analysed for this
to vessel strikes and entanglements in slack buoy lines.107,309,310 The assessment indicate interactions and entanglements with marine
risk of sharks interacting with offshore farms can be managed through shellfish longline gear and finfish cages worldwide pose some level of
site selection to avoid known aggregation areas of local predators, the risk for protected species in coastal and offshore environments (see
use of robust containment barriers, and continuous monitoring and Table 6 for a summary of stressors and mitigation measures). This
removal of dead or injured fish.9,26,289,299 study focused on validated, credible reports of interactions and entan-
glement. We acknowledge that global systems for aquaculture regula-
tion and resource conservation are highly varied and generally reflect
3.5 | Marine debris the socioeconomic development, industrialisation, and political
systems of a nation or region. For countries without regulations, over-
Similar to fishing gear, lost or discarded aquaculture gear from a facil- sight, and accountability, the extent of coastal and offshore aquacul-
ity can contribute to marine debris.312,313 Potential sources of marine ture interactions with protected species is unknown or it is believed
debris from aquaculture operations include rope, nets, buoys, feed that voluntary reports (self-reports) underestimate interactions. It is
bags, plastics, cardboard, wood, rags, tools, syringes, plastic bags, important to note that offshore aquaculture, aquaculture with expo-
paper waste, oil filters, scrap metal, and general human trash (wrap- sure to the weather and ocean climate, generally requires a level of
pers, cups, cans etc.). While it may be difficult to determine the origin technologically advanced infrastructure, sophistication, and invest-
of buoys, lines, or loose ropes, other debris, such as feedbags, may be ment available in developed countries. As a product of becoming a
more easily traced to aquaculture activities. Marine wildlife is developed country with a mature economy, these countries generally
impacted by marine debris through ingestion, entanglement, bioaccu- have established environmental policies to protect natural elements
mulation, and habitat effects119,314,315; and the relative contribution from agricultural and industrial processes. This study did not include
312
of aquaculture gear is uncertain. Fishing nets, line, rope, and other marine aquaculture in coastal environments that has been observed
debris entangles, disfigures, and drowns wildlife by encircling or to be rapidly expanding in many developing countries and has put
ensnaring the animals. Infection and debilitation could occur if animals enormous pressure on natural resources and environmental sustain-
get lacerations or other wounds from debris.314,316 A marine species' ability (e.g., Shandong Province, China—the world's most productive
mobility is reduced when it becomes entangled in debris. The animal region for farmed shellfish).322,323
may suffer from starvation, suffocation, exhaustion, and increased risk Marine aquaculture siting and sustainable development belongs
of predation due to restricted movement.314,317 Marine debris can be in an ecologically responsible framework, taking into consideration all
inhaled accidentally, but often animals feed on it because it resembles potential interactions and effects on vulnerable species and their habi-
318,319
their food. The materials may accumulate in the animal's stom- tats. Spatial planning and siting of farms to avoid and minimise inter-
ach and cause malnutrition or starvation once ingested. Sharp objects actions with populations of protected marine species is an effective
can damage the mouth, digestive tract, or stomach lining, resulting in strategy for minimising negative interactions.324 Our review of histori-
nutrition loss, infection, sickness, starvation, and even death.320,321 In cal cases around the world suggests that while it appears that aqua-
addition, ingested items may block air passages and cause suffocation. culture does not indicate significant impacts to marine mammals, sea
Consumption of some debris items may also result in the release of turtles, seabirds, and sharks from documented marine aquaculture
harmful chemicals. To prevent farm waste from becoming marine entanglement events, additional monitoring to verify is warranted. It
debris, waste management and accountability are imperative. Detailed is unclear if the low incidence is because farms are relatively benign
site waste management plans will provide specific instructions for the and pose little risk, or because the number and density of farms and
fate of supplies (i.e., ropes, netting, plastics, cardboard, paper, steel the detection level for harmful interactions are considerably low. Fur-
drums, chemical containers, scrap metals) to prevent or minimise thermore, low frequency entanglements of critically endangered spe-
waste production, control waste, and responsibly recycle, reuse, or cies can result in a significant impact.
dispose waste. Gear at abandoned and derelict farm sites poses a Improved information about home ranges, movements, and
threat to wildlife and habitats when left behind. Debris caused by cat- behaviours of protected marine species in response to aquaculture farm-
astrophic events (e.g., hurricanes or typhoons) that compromise the ing could help inform aquaculture development and provide better
structural integrity of farm gear pose a threat to marine mammals, sea understanding of risks to wildlife.325 Understanding non-lethal effects of
turtles, seabirds, and sharks. Detailed company recovery plans should disturbance on protected species physiology or behaviour and the
emphasise immediate remedial action and specify technology and potential population level repercussions is key to a comprehensive risk
resources that will be used to rapidly recover equipment. assessment and understanding cumulative effects.79,80,326,327 (Figure 4).
 Summary of marine aquaculture stressors, sources, and impacts on protected species, including potential solutions and mitigation measures.
22

TABLE 6

Stressors Sources Impacts Vulnerable species Potential solutions/mitigation measures Key references
Habitat Farm construction, Behavioural change, Cetaceans, pinnipeds, • Conduct a farm risk assessment prior to 26,36,43,55,57,58,60–
modification, operations, and modified access to turtles, seabirds, structures being placed in the water. Include 63,65,66,68,70,73,76–80,84–
exclusion, or maintenance foraging and sharks proximity to critical, sensitive, or protected 87,89,91,92,94–103,107–
competition for activities reproduction areas, species and habitats, as well as a description of 111,147,168,204,239,241,244,261–
space reduction in fitness, potential impacts on biodiversity. 265,268,269,271,274,289,290,292
death • Site farms in areas that minimise the likelihood of
overlap with the migration routes or critical
breeding and feeding habitats of protected
species. Locate farms away from haul-out sites
and rookeries.
• Reduce fish feed waste to minimise attraction to
farms and the time protected species and marine
life spend near pens.
• Log marine mammal presence and behaviour
around farms including night-time feeding activity
around lighted areas. Report incidents to the
appropriate authorities and keep records of
entanglements and mortalities.
Physical gear Buoy lines, anchor Behavioural change, Cetaceans, pinnipeds, • Minimise the number of vertical lines in the water 9,26,36,43,53,55,56,58–60,64,67,70–
interaction, lines, exclusion injury, death turtles, seabirds, to reduce entanglement risks. 72,78,84,87,88,105,128–
entanglement, or nets, cage mesh sharks • Enact a predator avoidance plan including gear 131,135–158,208,237,245,246,266,
entrapment maintenance and repair schedules. Keep all 272,280,281,289,290,299,309–311
anchor and backbone lines properly tensioned
and nets maintained (cleaning, repairing holes).
• Consider antipredation measures as preventative
and only second as a cure to problems.
• Exclusion measures including predator nets, top
nets, electric fences, and other deterrents should
be installed at the start of the farming operation.
This practice will prevent training potential
predators to locate a food source and develop
predatory behaviours around the farm.
• Enclosing predator nets at the bottom of fish
cages will prevent mammals and diving birds from
getting trapped and potentially drowning.
• Develop a communication plan and coordinate
with local wildlife authorities trained in rapid
animal rescue response in case an animal is
trapped or entangled in farm gear.
• Practice good husbandry techniques at fish farms
to ensure the health of the fish and prevent
predation. Remove mortalities from cages and
surrounding area and keep food sealed in
containers to avoid attracting wildlife to the site.
BATH ET AL.
TABLE 6 (Continued)

Stressors Sources Impacts Vulnerable species Potential solutions/mitigation measures Key references
BATH ET AL.

• Include a farm decommissioning strategy in the

operating plan to maximise efficient gear removal,
eliminate marine debris, and minimise
opportunities for animal entanglement.
Underwater noise Vessels, feeding Behavioural change, Cetaceans, pinnipeds, • Reduce vessel speed in the vicinity of animals. 26,43,68,175–180,182–
systems, trigger stress turtles • Minimise the noise levels at the source (e.g., 190,208,211,213,217,223
generators, response, feeding systems, generators, equipment at the
aerators, net communication lowest operable noise level).
cleaning disruption, disrupt • Separate the noise-producing activity spatially
equipment, foraging efficiency, and/or temporally from the sensitive species.
acoustic deterrents cause habitat • Avoid or minimise the noise-producing activity
displacement or within close proximity to sensitive habitats,
avoidance, defining appropriate clearance distances where
temporary or necessary.
permanent injury, • Assess the ambient soundscape environment pre-
death construction. Test, measure, and monitor noise
produced by equipment during construction,
operation, and maintenance periods. Compare
with the sound frequency and levels at which key
marine species produce and receive sound.
Practice adaptive management to minimise
adverse impacts to resident or transient species in
the farm vicinity.
Artificial light Lights for navigation, Behavioural change Cetaceans, pinnipeds, • Limit the use of underwater lighting. Only use 55,60,71,122,163,199–204,261–263,271
personal safety, turtles, seabirds, appropriate submerged lighting for beneficial
farm security, sharks outcomes.
submerged lights • Lights should be shielded from all but essential
for fish growth, directions. Spotlights should be positioned high
maturation above the water so penetration is maximised and
schedules, reduce reflection is minimised.
fish densities, and • Restrict lighting on moored vessels at night to the
evenly distribute minimum required for safe operation
fish in cages
Presence of vessels Required for Vessel collision Cetaceans, turtles, • Reduce number of vessels used during operations 191–195,255–257
construction, resulting in injury surface-dwelling and maintenance as possible.
operations, and and/or death, sharks (i.e., basking • Train vessel crew as lookouts to spot and identify
maintenance navigation hazard, and whale sharks) animals in the vicinity.
habitat • Operate vessels at slow speeds when performing
modification, work within and around the farm. Travel at
behavioural speeds necessary for safe and efficient navigation,
change, reduction i.e., at speeds necessary to maintain steerage if
in fitness towing equipment, but not so fast that objects in
the water cannot be avoided.
23

(Continues)
 (Continued)
24

TABLE 6

Stressors Sources Impacts Vulnerable species Potential solutions/mitigation measures Key references
• When animals are sighted, maintain a minimum
approach distance between the vessel and the
animal.
• Avoid excessive speed or abrupt changes in
direction until the animal has left the area.
• Reduce speed and shift the engine to neutral
when an animal is sighted in the vessel's path or
in close proximity to a moving vessel and when
safety permits. Do not engage the engines until
the animals are clear of the area.
• Traverse the same designated vessel transport
channels to and from the farm.
Marine debris Rope, nets, buoys, Entanglement or Cetaceans, pinnipeds, • Design a site waste management plan to prevent 119,312–321
feed bags, plastics, ingestion resulting turtles, seabirds, or minimise waste, control waste, and responsibly
cardboard, wood, in sickness, sharks recycle or dispose waste. This plan should include
rags, tools, starvation, injury, specific instructions for the fate of supplies (i.e.,
syringes, plastic infection, reduction ropes, netting, plastics, cardboard, paper, steel
bags, paper waste, in fitness, death drums, chemical containers, scrap metals).
oil filters, scrap Evaluate and renew plan annually.
metal, and general • Source uniquely manufactured farm gear or
human trash employ gear marking to allow the materials to be
tracked back to specific farms. For example, rope
designed with unique patterns can be used so
that it can be identified (and quantified) as
belonging to a certain farm if it is lost as marine
debris.
BATH ET AL.
BATH ET AL. 25

F I G U R E 4 “The Population Consequences of Disturbance (PCoD) conceptual framework, modified from National Academies.79 The boxes
within the dashed grey boundary line represent the effects of exposure to a stressor and a range of ecological drivers on the vital rates of an
individual animal. The effects are then integrated across all individuals in the population to project their effects on the population's dynamics”
(reproduced from Pirotta et al.80).

There is also a great need for understanding ecosystem shifts and research. Addressing the potential for protected species interactions
impacts on species abundance and distribution. For example, aquaculture with aquaculture infrastructure and gear, improved monitoring proto-
may influence species composition locally (e.g., attracting of predators) cols, event documentation, as well as mitigation strategies is impera-
thus influencing local movements of protected species around the farm. tive for responsible aquaculture growth in the ocean. This summary,
There are technological, monitoring, and siting approaches that including best farm management practices and monitoring, will glob-
may provide risk reduction for protected species interacting with aqua- ally inform industry planning, the regulatory processes, and robust
culture farms. In the United States, mitigation measures are commonly species conservation strategies.
explored during pre-application and permitting processes. Practices that
warrant more attention include adaptive monitoring, smart design using AUTHOR CONTRIBU TIONS
entanglement simulator technologies,328 use of breakaways and ten- Gretchen E. Bath: Investigation; writing – original draft; methodology;
sion sensors, and innovative deterrent technology. Animal morphology validation; writing – review and editing; formal analysis; project
and interactions with fishing gear can inform how mechanics and administration; data curation. Carol A. Price: Investigation; conceptu-
behaviours such as swimming speed, tail-beat frequency, water column alization; writing – review and editing. Kenneth L. Riley: Funding
placement, feeding position, veer away, roll, and startle reflex, increase acquisition; writing – original draft; writing – review and editing;
the probability of aquaculture gear entanglement. Identifying how ani- project administration; resources; supervision. James A. Morris:
mals use visual and acoustic senses to detect farm structures will Conceptualization; funding acquisition; writing – original draft; writing –
advance technical solutions to avoid interactions. Quantitative data on review and editing; project administration; resources; supervision.
the properties of mooring lines and cages such as tensile strength,
bending stiffness, elongation, friction, and wear due to internal and AC KNOW LEDG EME NT S
external damage define variables and parameters for developing Support for this work was provided by the NOAA Fisheries, Office of
models. Engineering advances and interaction simulation models can Aquaculture and the NOAA National Ocean Service, National Centers
improve aquaculture gear design to minimise entanglement events and for Coastal Ocean Science. The authors are grateful to the following
reduce injuries and fatalities. people (in alphabetical order) for sharing their expertise and their
This extensive review highlights interactions between aquacul- thorough reviews of this review paper: Dr. Tomma Barnes, Dr. Ann
ture farms and vulnerable species, underscores the potential risks Bowles, Langley Gace, Scott Lindell, David Morin, Penny Ruvelas, and
involved, and identifies critical areas of research to incite collaborative several reviewers in the NMFS Office of Protected Resources and
26 BATH ET AL.

Protected Species Divisions in the US regions. The authors appreciate Technologies in Aquaculture. Woodhead Publishing Ltd Press and
the insightful reviews provided by three anonymous experts who CRC Press; 2009:914-944.
16. Goudey CA. Practical aspects of offshore aquaculture system design.
helped improve the manuscript quality.
Paper presented at: World Aquaculture Society Aquaculture America;
February 17. 2009 Seattle, WA. https://docplayer.net/50002927-
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